HANDBOOK 

OF 

PHYSIOLOGY 


BLDDO- SPECTRA  CDMPARED  WITH  SPECTRUM  DP  ARGAND- LAMP 


1  Spectrum  oP  Ardand-lamp  with  PraunhoPErs  lines  in  position. 

2  Spectrum  oP  Dxyhsmodlobin  in  diluted  blood. 
.'3  Spectrum  aP  Reduced  nsmoQlobin. 

4  Spectrum  aP  Carbonic  oxide  Hsmp^iobio. 
o  Spectrum  oP  AcidHsmatm  in  ethErial  solution. 

6  Spectrum  oP  Alkaline  HaEmatin. 

7  Spectrum  Dp  Chloroform  extract  oT  acidulated  Qx-Bile. 

8  Spectrum  nP  MethffimaQlabin. 

9  Spectrum  oP  HasmachromDi^Gn. 
10  Spectrum  op  Hsmetopnrphyrin. 

Mosfofth?  above  SjM>ctnr  have  keen  (frown  fmrn  observations  by  MTWblpraik  F.C.S. 


KIRKES'   HANDBOOK 


OF 


PHYSIOLOGY 


Revised  and  Rewritten  by 

CHARLES  WILSON  GREENE,  A.  M.,  Ph.  D. 

PROFESSOR   OF   PHYSIOLOGY   AND   PHARMACOLOGY,    UNIVERSITY   OF   MISSOURI 


ftentb  American  IRcvtston 


WITH  FIVE  HUNDRED  AND  TWENTY-FOUR  ILLUSTRATIONS, 
INCLUDING  MANY  IN  COLORS 


: 


:2Z 


NEW   YORK 
WILLIAM  WOOD  AND  COMPANY 

MDCCCCXXII 


COPYRIGHT,  1922 
BY  WILLIAM  WOOD  &  COMPANY 


THE  MAPLE  PRESS  YORK  PA 


PREFACE 


In  this  the  tenth  American  edition  of  the  Handbook  of  Physiology 
revisions  and  amplifications  have  been  made  throughout  the  entire  text. 
New  subject  matter,  new  illustrations,  and  the  more  recent  refinements  of 
methods  have  been  freely  incorporated.  This  has  unavoidably  increased 
the  size  of  the  volume.  The  chapters  on  Circulation,  Respiration,  Internal 
Secretion,  Metabolism  and  the  Autonomic  division  of  the  Nervous  System 
in  particular  have  been  entirely  rewritten  and  reillustrated. 

The  chapter  on  the  Circulation  has  been  made  to  include  the  newer 
researches  on  the  development  of  cardiac  physiology  as  regards  rhythm 
production,  the  control  and  finer  adjustments  of  rate,  and  the  conduction 
phenomena  that  determine  sequence.  These  factors,  represented  in  the 
highly  differentiated  bundle  branch  system,  are  given  new  emphasis  and 
new  illustration.  Physiological  interest  in  Respiration  has  been  inten- 
sified by  the  practical  problems  of  air  navigation  and  by  the  newer  inves- 
tigations in  the  field  of  oxygen  supply  and  oxygen  control  in  relation  to  the 
daily  physiological  round  in  both  health  and  disease.  The  result  has 
been  a  new  impetus  to  respiratory  physiology  contributed  to  by  numerous 
writers  of  the  last  decade.  The  works  of  Barcroft,  Henderson,  Schneider, 
Greene  and  Gilbert  and  numerous  others  have  been  drawn  on  from  this 
field. 

The  science  of  nutrition  has  made  rapid  advances.  That  subject  has 
been  revised  to  call  attention  to  the  very  fundamental  work  of  Osborne 
and  Mendel  on  food  factors  necessary  for  growth;  of  Funk,  Voegtlin  and 
others  on  the  vitamines  and  nutritional  diseases;  of  Lusk  and  DuBois  in 
the  field  of  basal  metabolism;  of  Van  Slyke,  Stadie,  Harrop,  and  others 
on  blood  gases,  and  of  Banting,  Best  and  Macleod  on  glycemia  and  the 
hormone  of  the  pancreas  controlling  sugar  metabolism. 

The  giant  strides  of  the  science  of  physiology  make  it  difficult  for  a 
textbook  to  keep  pace  with  the  literature,  but  it  is  hoped  that  the  newest 
facts  and  principles  have  been  incorporated  in  so  far  as  the  limitations 
of  the  available  space  permit.  Many  of  the  illustrative  laboratory  experi- 
ments have  been  again  rewritten,"  and  improvements  simplifying  the 
experimental  technique  have  been  incorporated.  It  is  felt  that  the  student 
in  Physiology  gains  the  greatest  strength  in  laboratory  experience  when 
the  tests  he  executes  are  chosen  from  the  standpoint  of  the  efficiency  of  the 
entire  work.  In  this  field,  under  the  present  day  conditions,  the  deter- 


VI  PREFACE 

mining  pedagogical  horizon  includes  not  only  the  subject  matter  of 
Physiology,  but  of  Physics  and  Chemistry  on  which  Physiology  rests,  as 
well  as  Clinical  Medicine  and  Surgery  for  which  Physiology  furnishes  the 
foundation. 

For  valuable  aids  and  criticisms  in  this  and  recent  editions  I  again 
acknowledge  grateful  indebtedness  to  Professor  Robert  Banks  Gibson, 
University  of  Iowa,  Dr.  Carl  Hartley  Greene,  Mayo  Foundation,  to  my 
colleague,  Professor  Addison  Gulick,  and  to  my  students  and  assistants 
of  former  years,  Professor  Theodore  K.  Kruse  of  the  University  of 
Pittsburgh,  and  Professor  Erwin  Ellis  Nelson  of  the  University  of 
Michigan. 

CHAS.  W.  GREENE. 
COLUMBIA,  Mo., 
Sept.  i,  1922. 


CONTENTS 


PAGE 

CHAPTER  I — THE  PHENOMENA  or  LIFE;  Properties  of  Protoplasm, 

Structure  of  Protoplasm i 

CHAPTER  II — CELL  DIFFERENTIATION  AND  THE  STRUCTURE  OF  THE 
ELEMENTARY  TISSUES;  The  Structure  of  the  Cell,  The  Structure 
of  the  Elementary  Tissues.  The  Epithelial  Tissues.  The 
Connective  Tissues.  Muscular  Tissue.  Nervous  Tissue.  17 

CHAPTER  III— THE  CHEMICAL  COMPOSITION  OF  THE  BODY;  The 
Nitrogenous  Substances,  The  Proteins,  Classification  of  the  Pro- 
teins, Characteristics  of  the  Proteins,  The  Fats,  The  Carbo- 
hydrates; Inorganic  Substances  of  the  Body,  Laboratory  Experi- 
ments  79 

CHAPTER  IV— THE  BLOOD;  Quantity  of  the  Blood,  Coagulation  of 
the  Blood,  Morphology  of  the  Blood,  Chemical  Composition  of 
the  Blood,  Globulocidal  and  Other  Properties  of  Serum,  The 
Character  and  Composition  of  Lymph,  Laboratory  Experiments  117 

CHAPTER  V — THE  CIRCULATION  OF  THE  BLOOD;  Anatomical  Con- 
siderations, The  Action  of  the  Heart,  The  Regulative  Influence  of 
the  Central  Nervous  System,  The  Circulation  through  the  Blood- 
vessels, The  Pulse,  The  Peripheral  Regulation  of  the  Flow  of 
Blood,  Vaso-constrictor  and  Vaso-dilator  Nerves  for  Individual 
Organs,  Laboratory  Experiments 166 

CHAPTER  VI— RESPIRATION;  The  Respiratory  Apparatus,  The 
Movements  of  the  Respiratory  Mechanism,  Respiratory  Changes 
in  the  Air  Breathed,  The  Respiratory  Changes  in  the  Blood,  The 
Nervous  Regulation  of  the  Respiratory  Apparatus,  The  Effect  of 
Respiration  on  the  Circulation,  Laboratory  Experiments  in 
Respiration 278 

CHAPTER  VII — SECRETION  IN  GENERAL;  Organs  and  Tissues  of 
Secretion,  Secreting  Glands,  The  Process  of  Secretion,  Influence 
of  the  Nervous  System  on  Secretion 335 

vii 


Vlll  CONTENTS 

PAGE 

CHAPTER  VIII— FOOD  AND  DIGESTION;  Food  and  Food  Principles, 
The  Process  of  Digestion,  Digestion  in  the  Mouth,  Deglutition, 
Nervous  Mechanism  of  Deglutition,  Digestion  in  the  Stomach, 
Movements  of  the  Stomach,  Digestion  in  the  Intestines,  Move- 
ments of  the  Intestines,  Laboratory  Experiments  in  Digestion, 
Saliva  and  Salivary  Digestion,  Gastric  Juice  and  Gastric  Diges- 
tion, Pancreatic  Juice  and  Pancreatic  Digestion 341 

CHAPTER  IX — ABSORPTION;  Absorption  in  the  Stomach,  Absorp- 
tion in  the  Intestines,  Absorption  from  the  Skin ,  the  Lungs,  etc.  .  41 1 

CHAPTER  X— EXCRETION;  Structure  and  Function  of  the  Kidneys, 
General  Structure,  The  Urine,  The  Method  of  Excretion  of  Urine, 
The  Discharge  of  the  Urine,  The  Structure  and  Excretory  Func- 
tions of  the  Skin,  Laboratory  Experiments  in  Excretion  .  .  .  .421 

CHAPTER  XI — METABOLISM,  NUTRITION,  AND  DIET;  Metabolism  of 
Proteids,  The  Metabolism  of  Fats,  the  Metabolism  of  Carbohy- 
drates, Influence  of  Minerals,  Fasting,  Requisites  of  a  Normal 
Diet,  The  Vitamines,  The  Influence  of  the  Ductless  Glands  on 
Metabolism 454 

CHAPTER  XII — ANIMAL  HEAT;  Heat-producing  Organs,  Variation 
in  the  Loss  of  Heat,  Variation  in  the  Production  of  Heat,  Influ- 
ence of  the  Nervous  System  on  Heat  Production 501 

CHAPTER  XIII — MUSCLE-NERVE  PHYSIOLOGY;  Chemical  Composi- 
tion of  Muscle,  The  Properties  of  Living  Muscle,  Single  Muscle 
Contractions,  Conditions  which  Affect  the  Irritability  of  the 
Muscle  and  the  Character  of  the  Contraction,  Tetanic  and  Volun- 
tary Muscular  Contractions,  The  Type  of  Contraction  in  Invol- 
untary Muscle  and  in  Cilia,  The  Function  of  Nerve  Fiber,  Some 
Special  Coordinated  Motor  Activities,  Locomotion,  The  Produc- 
tion of  the  Voice,  Laboratory  Experiments  on  Muscle  and  Nerves.  5 10 

CHAPTER  XIV— THE  NERVOUS  SYSTEM;  Function  of  the  Nerve  Cell; 
Specific  Energy  of  the  Nerve  Impulse,  Structure  and  the  Function 
of  the  Spinal  Cord,  Tracts  of  the  Cord,  The  Functions  of  the  Cord. 
The  Brain,  The  Medulla  Oblongata  and  Pons,  Structure,  Func- 
tions of  the  Medulla,  The  Cerebellum,  The  Midbrain,  The 
Peduncles  of  the  Cerebrum,  Corpora  Quadrigeminaj  Corpora 
Genic.ulata,  Corpora  Striata,  The  Cerebrum,  Structure  of  the 
Cortex,  General  Function  of  the  Cerebrum,  Localization  of  the 


CONTENTS  ix 

PAGE 

Motor  Function,  Localization  of  Sensory  Function,  Association 
Centers,  The  Cranial  Nerves,  The  Sympathetic  or  Autonomic 
System,  The  Physiology  of  Sleep,  Laboratory  Experiments  on 
the  Nervous  System 572 

CHAPTER  XV— THE  SENSES;  I.  The  Senses  of  Touch,  Pain,  Tem- 
perature, and  the  Muscle  Sense.  II.  Taste  and  Smell,  The  Sense 
of  Taste,  The  Sense  of  Smell.  III.  Hearing  and  Equilibration, 
The  Anatomy  of  the  Ear,  The  Physiology  of  Hearing,  The  Sense 
of  Equilibrium.  IV.  The  Sense  of  Sight,  The  Eye,  The  Optical 
Apparatus,  Accommodation,  Defects  in  the  Optical  Apparatus, 
Visual  Sensations  from  Excitation  of  the  Retina,  Color  Sensations, 
Binocular  Vision,  Visual  Judgments,  Laboratory  Directions  for 
Experiments  on  the  Sense  Organs 679 

CHAPTER  XVI— THE  REPRODUCTIVE  ORGANS;  The  Reproductive 
Organs  of  the  Male,  The  Reproductive  Organs  of  the  Female, 
Ovulation  and  Menstruation,  Menstrual  Life 768 

CHAPTER  XVII— DEVELOPMENT;  Changes  which  Occur  in  the 
Ovum  Prior  to  Impregnation,  Changes  Following  Impregnation, 
Circulation  of  Blood  in  the  Fetus,  Parturition,  Lactation  .  .  .  781 

INDEX 797 


HANDBOOK  OF  PHYSIOLOGY 


CHAPTER  I 
THE  PHENOMENA  OF  LIFE 

PHYSIOLOGY  is  the  science  which  treats  of  the  various  processes  or  changes 
which  take  place  in  the  organs  and  tissues  of  the  body  during  life.  These 
processes,  however,  must  not  be  considered  as  by  any  means  peculiar  to  the 
human  organism,  since,  putting  aside  the  properties  which  serve  to  distin- 
guish man  from  other  animals,  the  changes  which  go  on  in  the  tissues  of  man 
go  on  in  much  the  same  way  in  the  tissues  of  all  other  animals  as  long  as  they 
live.  Furthermore,  it  is  found  that  similar  changes  proceed  in  all  living 
vegetable  tissues;  they  indeed  constitute  what  are  called  vital  phenomena, 
and  are  those  properties  which  mark  out  living  from  non-living  material. 

The  lowest  types  of  life,  whether  animal  or  vegetable,  are  found  to  con- 
sist of  minute  masses  of  a  substance  generally  known  under  the  name  of 
protoplasm.  Each  such  living  mass  is  called  a  cell,  so  that  these  minute 
elementary  organisms  are  designated  unicellular. 

The  phenomena  of  life  are  exhibited  by  protoplasm,  whether  that  exists 
in  the  simple  form  typified  by  a  microscopic  one-celled  animal,  or  in  a  more 
complex  mass  represented  by  the  organs  and  tissues  of  animals  and  plants. 
In  the  lowest  type  of  life  the  morphological  unit  of  structural  organization  is 
represented  by  the  single  cell.  In  the  more  complex  organisms  of  both  ani- 
mals and  plants  the  total  mass  represents  a  great  aggregation  of  more  or  less 
distinct  cells.  A  degree  of  differentiation  takes  place  whereby  the  tissues 
and  organs  of  the  body  of  plants  and  animals  present  great  aggregates  of 
differentiating  cells.  It  must  be  at  once  evident  that  the  great  mass  of  knowl- 
edge dealing  with  the  nature  and  activities  of  protoplasm  constitutes  the 
science  of  physiology.  The  cell,  therefore,  is  the  working  unit  in  physiology 
no  less  than  in  morphology. 

The  prime  importance  of  the  cell  as  an  element  of  structure  was  first 
established  by  the  researches  of  the  botanist  Schleiden,  and  his  conclusions, 
drawn  from  the  study  of  vegetable  histology,  were  at  once  extended  by  Theo- 
dor  Schwann  to  the  animal  kingdom.  The  earlier  observers  defined  a  cell 
as  a  more  or  less  spherical  body  limited  by  a  membrane,  and  containing  a 
smaller  body  termed  a  nucleus,  which  in  its  turn  incloses  one  or  more  still 

I 


THE    PHENOMENA    OF   LIFE 


Space  contain- 
ing liquid. 


Protoplasm. 


——Nucleus. 


—  Cell  wall. 


FIG.  i. — Vegetable  Cells. 


smaller  bodies  or  nucleoli.  Such  a  definition  applied  admirably  to  most  vege- 
table cells,  but  the  more  extended  investigation  of  animal  tissues  soon 
showed  that  in  many  cases  no  limiting  membrane  or  cell  wall  could  be 
demonstrated. 

The  presence  or  absence  of  a  cell  wall, 
therefore,  was  then  regarded  as  quite  a 
secondary  matter,  while  at  the  same  time  the 
cell  substance  came  gradually  to  be  recog- 
nized as  of  primary  importance.  Many  of 
the  lower  forms  of  animal  life,  the  Rhizopoda, 
were  found  to  consist  almost  entirely  of 
matter  very  similar  in  appearance  and  chem- 
ical composition  to  the  cell  substance  of 
higher  forms;  and  this  from  its  chemical 
resemblance  to  flesh  was  termed  Sarcode 
by  Dujardin.  When  recognized  in  vege- 
table cells  it  was  called  Protoplasm  by 
Mulder,  while  Remak  applied  the  same  name 
to  the  substance  of  animal  cells.  As  the 
presumed  formative  matter  in  animal  tissues 

it  was  termed  Blastema,  and  in  the  belief  that,  wherever  found,  it  alone  of 
all  substances  has  to  do  with  generation  and  nutrition,  Beale  has  named 
it  Germinal  matter  or  Bioplasm.  Of  these  terms  the  one  most  in  use 
at  the  present  day  as  we  have  already  said,  is  protoplasm,  and  inasmuch  as 
all  life,  both  in  the  animal  and  vegetable  kingdoms,  is  associated  with 
protoplasm,  we  are  justified 
in  describing  it,  with  Huxley, 

i        <k    i        -11        •        P   IT     )>  /^i^S^SwSSBSSS^ilK  Nucleus   or    ger- 

as   the  "physical  basis  of  life,"       .K^BV-- —   minai  vesicle. 

••«!••  » 

or  simply  "living  matter." 

General  Physical  a  n  d 
Chemical  Properties  of  Pro- 
toplasm. — Protoplasm  is  a 
semifluid  substance,  which  ab- 
sorbs, but  does  not  mix  with 
water.  It  is  transparent  and 

generally  colorless,  with   refrac-  FlG-  2.— Semidiagrammatic  Representation  of 

,    a  Human  Ovum,  showing  the  parts  of  an  animal 
tive  index  higher  than   that  of  ce\\m    (Cadia.) 

water,  but  lower  than  that  of  oil. 

It  is  neutral  or  weakly  alkaline  in  reaction,  but  may  under  special  cir- 
cumstances be  acid,  as,  for  example,  after  activity.  It  undergoes  heat 
coagulation  at  a  temperature  of  about  54.5°  C.  (130°  F.),  and  hence  no 
organism  can  live  when  its  own  temperature  is  raised  above  that  point. 
It  is  also  coagulated  and  therefore  killed  by  alcohol,  by  solutions  of 


minal  spot. 
----Space  left  by  re- 
traction of  yolk. 

—  Vitellus  of  yolk. 


Vitelline    mem- 
brane. 


CHARACTERISTICS  OF  PROTOPLASM  3 

many  of  the  metallic  salts,  by  strong  acids  and  alkalies,  and  by  many 
other  chemical  substances. 

Under  the  microscope  it  is  seen  almost  universally  to  be  granular,  the 
granules  consisting  of  different  substances,  albuminous,  fatty,  or  carbo- 
hydrate matter.  The  granules  are  not  equally  distributed  throughout  the 
whole  cell  mass,  as  they  are  sometimes  absent  from  the  outer  part  or  layer 
and  very  numerous  in  the  interior.  In  addition  to  granules,  protoplasm 
generally  exhibits  spaces  or  vacuoles,  usually  globular  in  shape,  except- 
ing during  movement,  when  they  may  be  irregular,  and  filled  with  a  watery 
fluid.  These  vacuoles  are  more  numerous  and  pronounced  in  vegetable 
than  in  animal  cells.  Gas  bubbles  also  sometimes  exist  in  cells. 

It  is  impossible  to  make  any  definite  statement  as  to  the  exact  chemical 
composition  of  living  protoplasm,  since  the  methods  of  chemical  analysis 


FIG.  3. — Phases  of  Ameboid  Movement. 

necessarily  imply  the  death  of  the  cell;  it  is  stated,  however,  that  protoplasm 
contains  75  to  85  per  cent,  of  water,  and  of  the  15  to  25  per  cent,  of  solids  the 
most  important  part  belongs  to  the  class  of  substances  called  proteins  or  al- 
bumins. Proteins  contain  the  chemical  elements  carbon,  hydrogen,  nitrogen, 
oxygen,  sulphur,  and  phosphorus,  the  last  two  in  very  small  quantities  only. 
A  protein-like  substance,  nudein,  found  in  the  nuclei  of  cells,  contains  phos- 
phorus in  greater  abundance.  In  the  cell  nucleus  a  compound  of  nuclein 
with  protein,  called  nucleoprotein,  forms  the  most  abundant  protein  sub- 
stance. Other  bodies  are  frequently  found  associated  with  the  proteins,  such 
as  glycogen,  starch,  cellulose,  which  contain  the  elements  carbon,  hydrogen, 
and  oxygen,  the  last  two  in  the  proportion  to  form  water,  and  hence  are 
termed  carbohydrates;  fatty  bodies,  containing  carbon,  hydrogen,  and  oxygen, 
but  not  in  proportion  to  form  water;  lecithin,  a  complicated  fatty  body  con- 
taining phosphorus;  cholesterin,  a  monatomic  alcohol;  chlorophyll,  the  color- 
ing matter  of  plants;  hemoglobin,  the  complex  animal  pigment;  inorganic 
salts,  particularly  the  chlorides  and  phosphates  of  calcium,  sodium,  and  potas- 
sium ;  ferments,  and  many  special  substances. 

The  General  Physiological  Characteristics  of  Protoplasm. — The 
properties  of  protoplasm  may  be  well  studied  in  the  microscopic  animal  called 
the  ameba,  a  unicellular  organism  found  chiefly  in  fresh  water.  These 
properties  may  be  conveniently  studied  under  the  following  heads: 

The  Power  of  Spontaneous  Movement. — When  an  ameba  is  observed 
with  a  high  power  of  the  microscope,  it  is  found  to  consist  of  an  irregular  mass 


THE   PHENOMENA    OF    LIFE 


of  protoplasm  containing  one  or  more  nuclei,  the  protoplasm  itself  being 
more  or  less  granular  and  vacuolated.  If  watched  for  a  minute  or  two,  an 
irregular  projection  is  seen  to  be  gradually  thrust  out  from  the  main  body; 
other  masses  are  then  protruded  until  gradually  the  whole  protoplasmic  sub- 


l. 


2. 


FIG.  4. — Changes  of  Form  of  a  White  Corpuscle,  Sketched  at  Brief  Intervals.     The  figures 
show  also  the  ingestion  of  two  small  granules.     (Schafer.) 

stance  is,  as  it  were,  drawn  over  to  a  new  position,  and  when  this  is  repeated 
several  times  we  have  locomotion  in  a  definite  direction,  together  with  a  con- 
tinual change  of  form.  These  movements,  figures  3  and  4,  are  observed  in 
such  cells  as  the  colorless  blood  corpuscles  of  higher  animals,  in  the  branched 
corneal  cells  of  the  frog  and  elsewhere,  and  are  termed  ameboid. 


FIG.  5. — Cell  of  Tradescantia  Drawn  at  Successive  Intervals  of  Two  Minutes. — The 
cell  contents  consist  of  a  central  mass  connected  by  many  irregular  processes  to  a  peripheral 
film,  the  whole  forming  a  vacuolated  mass  of  protoplasm,  which  is  continually  changing 
its  shape.  (Schofield.) 

The  remarkable  movement  of  pigment  granules  observed  in  the  branched 
pigment  cells  of  the  frog's  skin  by  Lister  are  also  probably  due  to  ameboid 
movement.  These  granules  are  seen  at  one  time  distributed  uniformly 
through  the  body  and  branched  processes  of  the  cell,  while  at  another  time 
they  collect  in  the  central  mass  leaving  the  branches  quite  colorless. 

This  movement  within  the  pigment  cells  might  also  be  considered  an  ex- 
ample of  the  so-called  streaming  movement  not  infrequently  seen  in  certain 
of  the  protozoa,  in  which  the  mass  of  protoplasm  extends  long  and  fine  proc- 


CHARACTERISTICS    OF    PROTOPLASM  5 

esses,  themselves  very  little  movable,  but  upon  the  surface  of  which  freely 
moving  or  streaming  granules  are  seen.  A  gliding  movement  has  also  been 
noticed  in  certain  animal  cells;  the  motile  part  of  the  cell  being  composed  of 
protoplasm  bounding  a  central  and  more  compact  mass.  By  means  of  the 
free  movement  of  this  layer,  the  cell  may  be  observed  to  move  along. 

In  vegetable  cells  the  protoplasmic  movement  can  be  well  seen  in  the 
hairs  of  the  stinging-nettle  and  Tradescantia  and  in  the  cells  of  Vallisneria. 
It  is  marked  by  the  movement  of  the  granules  nearly  always  embedded  in  it. 
For  example,  if  part  of  a  hair  of  Tradescantia,  figures  5  and  6,  be  viewed 
under  a  high  magnifying  power,  streams  of  protoplasm  containing  crowds 
of  granules  hurrying  along,  like  the  foot  passengers  in  a  busy  street,  are 
seen  flowing  steadily  in  definite  directions,  some  coursing  round  the  film 
which  lines  the  interior  of  the  cell  wall,  and  others  flowing  toward  or  away 
from  the  irregular  mass  in  the  center  of  the  cell  cavity.  Many  of  these  streams 
of  protoplasm  run  together  into  larger  ones  and  are  lost  in  the  central  mass, 
and  thus  ceaseless  variations  of  form  are  produced.  The  movement  of 
the  protoplasmic  granules  to  or  from  the  periphery  is  sometimes  called 
vegetable  circulation,  whereas  the  movement  of  the  protoplasm  round  the 
interior  of  the  cell  is  called  rotation. 

The  first  account  of  the  movement  of  protoplasm  was  given  by  Rosel  in 
1755,  as  occurring  in  a  small  Proteus,  probably  a  large  fresh- water  ameba. 
His  description  was  followed  twenty  years  later  by  Corti's  demonstration  of 
the  rotation  of  the  cell  sap  in  characeae,  and  in  the  earlier  part  of  the  last 
century  by  Meyer  in  Vallisneria,  1827;  Robert  Brown,  1831,  in  "Staminal 
Hairs  of  Tradescantia."  Then  came  Dujardin's  description  of  the  granular 
streaming  in  the  pseudopodia  of  Rhizopods  and  movements  in  other  cells  of 
animal  protoplasm  (Planarian  eggs,  von  Siebold,  1841;  colorless  blood 
corpuscles,  Wharton  Jones,  1846). 

The  Power  of  Response  to  Stimuli,  or  Irritability. — Although  the  move- 
ments of  the  ameba  have  been  described  above  as  spontaneous,  yet  they 
may  be  increased  under  the  action  of  external  agencies  which  excite  them 
and  are  therefore  called  stimuli.  If  the  movement  has  ceased  for  the  time, 
as  is  the  case  if  the  temperature  is  lowered  beyond  a  certain  point,  move- 
ment may  be  set  up  again  by  raising  the  temperature.  Contact  with  foreign 
bodies,  gentle  pressure,  certain  salts,  and  electricity  produce  or  increase  the 
movement  in  the  ameba.  The  protoplasm  is,  therefore,  sensitive  or  irritable 
to  stimuli,  and  shows  its  irritability  by  movement  or  contraction  of  its 
mass. 

The  effects  of  some  of  these  stimuli  may  be  thus  further  detailed: 

a.  Changes  of  Temperature. — Moderate  heat  acts  as  a  stimulant;  the 
movement  stops  below  o°  C.  (32°  F.),  and  above  40°  C.  (104°  F.);  between 
these  two  points  the  movements  increase  in  activity;  the  optimum  tempera- 


THE   PHENOMENA    OP   LIFE 


ture  is  about  37°  to  38°  C.  Exposure  to  a  temperature  even  below  o°  C. 
stops  the  movement  of  protoplasm,  but  does  not  prevent  its  reappearance  if 
the  temperature  is  raised;  on  the  other  hand,  prolonged  exposure  to  a  tem- 
perature of  a  little  over  40°  C.  kills  the  protoplasm  and  causes  it  to  enter 
into  a  condition  of  coagulation  or  heat  rigor. 

b.  Mechanical  Stimuli. — When  gently  squeezed  between  a  cover  and 
object-glass  under  proper  conditions,  a  colorless  blood  corpuscle  contracts 
and  ceases  its  ameboid  movement. 

c.  Nerve  Influence. — By  stimulation  of  the  nerves  of  the  frog's  cornea, 
contraction  of  certain  of  its  branched  cells  has  been  produced. 

d.  Chemical  Stimuli. — Water  generally  stops  ameboid  movement,  and  by 
imbibition  causes  great  swelling  and  finally  bursting  of  the  cells.     In  some 
cases,  however  (myxomycetes),  protoplasm  can  be  almost  entirely  dried  up, 
but  remains  capable  of  renewing  its  movements  when  again  moistened. 
Dilute  salt  solution  and  many  dilute  acids  and  alkalies  stimulate  the  move- 
ments temporarily.     Strong  acids  or  alkalies  permanently  stop  the  move- 
ments; ether,  chloroform,  veratrum,  and  quinine  also  stop  it  for  a  time. 

Movement  is  suspended  in  an  atmosphere  of  hydrogen  or  carbonic  acid 
and  resumed  on  the  admission  of  air  or  oxygen,  but  complete  withdrawal  of 
oxygen  will  after  a  time  kill  the  protoplasm. 

e.  Electrical. — Weak  currents  stimulate  protoplasmic  movement,  while 

strong  currents  cause  the  cells  to  assume   a 
spherical  form  and  to  become  motionless. 

The  Power  of  Digestion,  Respiration,  and 
Nutrition. — This  consists  in  the  power  which 
is  possessed  by  the  ameba  and  similar  animal 
cells  of  taking  in  food,  modifying  it,  building 
up  tissue  by  assimilating  it,  and  rejecting  what 
is  not  assimilated.  These  various  processes 
are  effected  in  some  one-celled  animals  by  the 
protoplasm  simply  flowing  around  and  en- 
closing within  itself  minute  organisms  such  as 
diatoms  and  the  like.  From  these  it  extracts 
what  it  requires,  and  then  rejects  or  excretes  the 
remainder,  which  has  never  formed  part  of  the 
body.  This  latter  proceeding  is  done  by  the 
cell  withdrawing  itself  from  the  material  to  be 
excreted.  The  assimilation  constantly  taking 
place  in  the  body  of  the  ameba  is  for  the 
purpose  of  replacing  waste  of  its  tissue  conse- 
quent upon  manifestation  of  energy.  The 
respiratory  process  of  absorbing  oxygen  goes 
on  at  the  same  time. 


FIG.  6. — Cells  from  the  Stam- 
inal  Hairs  of  Tradescantia.  A, 
Fresh  in  water;  J5,  the  same  cell 
after  slight  electrical  stimula- 
tion; a,  b,  region  stimulation; 
c,  d,  clumps  and  knobs  of  con- 
tracted protoplasm.  (Kuhne.) 


CHARACTERISTICS    OF    PROTOPLASM  7 

The  processes  which  take  place  in  cells,  both  animal  and  vegetable,  are 
summed  up  under  the  term  metabolism  (from  /xera/itoA.^,  change).  The 
changes  which  go  on  are  of  two  kinds,  viz.,  assimilation,  or  building  up,  and 
disassimilation,  or  breaking  down;  they  may  be  also  called,  using  the  nomen- 
clature of  Gaskell,  anabolism  or  constructive  metabolism,  and  catabolism  or 
destructive  metabolism.  In  the  direction  of  anabolism  two  processes  occur, 
viz.,  the  building  up  of  special  though  non-living  substances  from  materials 
which  it  takes  in,  and  secondly,  the  building  up  of  its  own  living  substance 
from  those  or  other  materials.  As  we  shall  see  in  a  subsequent  paragraph, 
the  process  of  'anabolism  differs  to  some  extent  in  vegetable  and  animal 
cells.  The  catabolism  of  the  cell  consists  in  the  disintegrative  chemical 
changes  which  occur  in  the  cell  substance  itself  or  in  substances  in  contact 
with  it. 

The  destructive  metabolism  of  a  cell  is  increased  by  its  activity,  but  goes 
on  also  during  quiescence.  It  is  probably  of  the  nature  of  oxidation,  and  re- 
sults in  the  evolution  of  carbon  dioxide  and  water  on  the  one  hand,  and  in  the 
formation  of  various  more  complex  chemical  substances  on  the  other,  some  of 
which  may  be  stored  up  in  the  cell  for  future  use,  and  are  called  secretions, 
and  others,  like  carbon  dioxide,  for  example,  and  bodies  containing  nitrogen, 
are  eliminated  as  excretions. 

The  Power  of  Growth. — In  protoplasm  it  is  seen  that  the  two  processes  of 
waste  and  repair  go  on  side  by  side,  and  so  long  as  they  are  equal  the  size 
of  the  animal  remains  stationary.  If,  however,  the  building  up  exceed  the 


FIG.  7. — Diagram  of  an  Ovum  (a)  Undergoing  Segmentation.  In  (b)  it  has  divided 
into  two,  in  (c)  into  four;  and  in  (d)  the  process  has  ended  in  the  production  of  the  so-called 
"mulberry  mass."  (Frey.) 

waste,  then  the  animal  grows;  if  the  waste  exceeds  the  repair,  the  animal 
wastes;  and  if  decay  goes  on  beyond  a  certain  point,  life  becomes  impossible 
and  the  animal  dies. 

The  power  of  increasing  in  size,  although  essential  to  our  idea  of  life,  is 
not,  it  must  be  recollected,  confined  to  living  beings.  A  crystal  of  common 
salt,  for  example,  if  placed  under  appropriate  conditions  for  obtaining  fresh 
material,  will  increase  in  size  in  a  fashion  as  definitely  characteristic  and  as 
easily  to  be  foretold  as  that  of  a  living  creature;  but  the  growth  of  a  crystal 
takes  place  merely  by  additions  to  its  outside;  the  new  matter  is  laid  on  par- 
ticle by  particle,  and  layer  by  layer,  and,  when  once  laid  on,  it  remains  un- 
changed. In  a  living  structure,  where  growth  occurs,  it  is  by  addition  of 
new  matter,  not  to  the  surface  only,  but  throughout  every  part  of  the  mass, 
and  this  matter  becomes  an  intimate  part  of  the  living  substance. 


s 


THE    PHENOMENA    OF    LIFE 


The  Power  of  Reproduction. — The  ameba,  to  return  to  our  former  illus- 
tration, when  the  growth  of  its  protoplasm  has  reached  a  certain  point,  mani- 
fests the  power  of  reproduction,  by  splitting  up  into  (or  in  some  other  way 
producing)  two  or  more  parts,  each  of  which  is  capable  of  independent 
existence.  The  new  amebae  manifest  the  same  properties  as  the  parent, 
perform  the  same  functions,  grow  and  reproduce  in  their  turn.  This  cycle 
of  life  is  being  continually  passed  through. 

In  more  complicated  structures  than  the  ameba,  the  life  of  individual 
protoplasmic  cells  is  probably  very  short  in  comparison  with  that  of  the  organ- 
ism they  compose;  and  their  constant  decay  and  death  necessitate  constant 
reproduction.  The  manner  in  which  this  takes  place  has  long  been  the  sub- 
ject of  investigation. 

The  exact  manner  of  reproduction  and  growth  of  protoplasm  is  a  matter 
of  great  complexity.  Those  who  have  already  learned  the  embryological 
story  have  the  foundations  laid  for  the  physiological  uses  made  of  this  mate- 


Cell  membrane. 


Cell  reticulum.  — -  ^ 


Membrane  of  nucleus. 


Achromatic  substance  of 

nucleus. 
Chromatic  substance  of 

nucleus. 


FIG.  8. — Cell  with  its  Reticulum  Disposed  Radially;  from  the  intestinal  epithelium  of  a 

worm.     (Carnoy.) 

rial.  Certain  of  the  essential  facts  are  set  forth  in  Chapter  II  a  little  later,  in 
so  far  as  the  reproduction  of  the  cellular  unit  is  concerned.  The  reproduction 
of  the  animal  organism  as  a  whole  is  a  still  more  complicated  story  and  is 
presented  in  the  last  chapter  of  this  volume. 


THE  DIFFERENTIATION  OF  PROTOPLASM 

The  Morphological  Unit. — Protoplasm  was  formerly  thought  to  be  homo- 
geneous. It  is  found,  however,  that  every  mass  of  living  protoplasm  has  one 
or  more  special  structures  imbedded  in  the  mass,  the  nuclei.  In  most  tissues 
of  the  mammalian  body  each  mass  of  protoplasm  is  more  or  less  distinctly 
subdivided  into  elemental  divisions,  corresponding  to  the  number  of  nuclei, 
the  cells.  The  cells  present  in  the  mass,  therefore,  represent  the  morphologi- 
cal units.  The  arrangements  of  these  units  as  to  size  and  space  relations 
constitute  the  form  and  mass  characteristics  of  the  tissues  on  the  one  hand, 
and  of  the  organs  on  the  other.  Where  there  is  no  distinct  marking  off  of 


STRUCTURE    OF    PROTOPLASM  9. 

the  mass  of  protoplasm  into  individual  cellular  units,  there  is  formed  what 
is  known  as  a  syncytium.  All  syncytial  structures  are  multi-nuclear,  other- 
wise their  protoplasm  is  not  unlike  that  of  the  cell  as  a  unit. 

The  fuller  detail  of  cellular  types  and  cell  structure  is  given  a  little  later 
in  Chapter  II.  However,  for  the  purpose  of  reference  to  this  point,  one  may 
here  call  attention  to  the  fact  that  the  principal  differentiations  of  the  proto- 
plasm of  the  cell  are  the  nuclei  and  the  cytoplasm.  The  cytoplasm  is  differ- 
entiated further  into  two  substances,  spongioplasm  and  hyaloplasm.  The 
spongioplasm  or  reticulum  forms  a  fine  network,  increases  in  relative  amount 
as  the  cell  grows  older,  and  has  an  affinity  for  staining  reagents.  The 
hyaloplasm  is  less  refractile,  elastic,  or  extensile,  and  has  little  or  no  affinity 
for  stains;  it  predominates  in  young  cells,  is  thought  to  be  fluid,  and  fills  the 
interspaces  of  the  reticulum.  The  nodal  points  of  the  reticulum,  with  the 
granular  microsomes,  found  in  the  protoplasm,  cause  the  granular  appear- 
ance. The  arrangement  of  the  reticulum  varies  considerably  in  different 
cells,  and  even  in  different  parts  of  the  same  cell. 

In  some  cells,  particularly  in  plants,  but  also  in  some  animal  cells,  there 
is  a  tendency  toward  the  formation  of  a  firmer  external  envelope,  constituting 
in  vegetable  cells  a  membrane  distinct  from  the  more  central  and  more  fluid 
part  of  the  protoplasm.  In  such  cases  the  reticulum  at  the  periphery  of 
the  cell  is  made  up  of  very  fine  meshes.  The  membrane  when  formed  is 
usually  pierced  with  pores  by  which  fluid  may  pass  in,  or  through  which 
protrusion  of  the  protoplasmic  filaments  forming  the  cell's  connection  with 
other  cells  surrounding  it  may  take  place. 


FIG.  9. — A:  The  Colorless  Blood  Corpuscle,  Showing  the  Intracellular  Network,  and 
two  nuclei  with  intranuclear  network.  B:  Colored  blood  corpuscle  of  newt  showing  the 
intracellular  network  of  fibrils.  Also  oval  nucleus  composed  of  limiting  membrane  and 
fine  intranuclear  network  of  fibrils.  X  800.  (Klein  and  Noble  Smith.) 

All  protoplasm  at  some  period  of  its  existence  possesses  one  or  more  neclei. 
The  origin  of  a  nucleus  in  a  cell  is  the  first  sign  of  the  differentiation  of  proto- 
plasm. The  existence  of  nuclei  was  first  pointed  out  in  the  year  1833  by 
Robert  Brown,  who  observed  them  in  vegetable  cells.  They  are  either  small 
transparent  vesicular  bodies  containing  one  or  more  smaller  particles  called 
nucleoli,  and  always  when  in  the  resting  condition  bounded  by  a  well-defined 


10  THE    PHENOMENA    OF    LIFE 

envelope.  In  their  relation  to  the  life  of  the  cell  they  are  certainly  hardly 
second  in  importance  to  the  cytoplasm  itself,  and  thus  Beale  is  fully  justified 
in  comprising  both  under  the  term  "germinal  matter."  The  nuclei  control 
the  nutrition  of  the  cell,  and  probably  initiate  the  process  of  subdivision.  If 
a  cell  be  mechanically  divided  so  that  a  portion  of  it  possesses  the  nucleus 
while  other  portions  have  no  nucleus,  that  portion  containing  the  nucleus  will 
live  and  develop  while  the  parts  without  nucleus  soon  die.  Concerning  this 
interesting  question  of  the  relation  of  the  nuclei  and  cytoplasm  in  the  cells, 
Schafer  summarizes  as  follows:  "There  are  cells  and  unicellular  organisms 
both  animal  and  vegetable,  in  which  no  reticular  structure  can  be  made  out, 
and  these  may  be  formed  of  hyaloplasm  alone.  In  that  case,  this  must  be 
looked  upon  as  the  essential  part  of  protoplasm.  So  far  as  ameboid  phenom- 
ena are  concerned  it  is  certainly  so;  but  whether  the  chemical  changes  which 
occur  in  many  cells  are  effected  by  this  or  by  spongioplasm  is  another  matter." 

Protoplasmic  nuclei  are  highly  differentiated  chemically  as  well  as  func- 
tionally. They  contain  special  structures  which  react  in  a  characteristic 
chemical  way  to  staining  solutions,  and  to  other  chemical  treatment.  This 
differential  structure  is  emphasized  in  Chapter  II.  There  the  morphological 
changes  through  which  the  nuclei  pass  in  cell  multiplication  are  given  in 
greater  detail.  It  is  the  study  of  these  changes  that  supplies  the  basis  of  fact 
for  many  of  our  present  conceptions  of  the  physiological  importance  of 
nuclei. 

Differentiation  and  Growth  of  Organized  Protoplasm. — The  detail  of  cellu- 
lar division  of  protoplasm  is  more  fully  given  in  Chapter  II.  The  morphologi- 
cal fact  to  which  attention  is  called  here  is  that  as  we  proceed  upward  in  the 
scale  of  life  from  the  unicellular  organisms,  another  phenomenon  is  exhibited 
in  the  life  history  of  the  higher  forms,  namely,  that  of  development.  The 
one-celled  ameba  comes  into  being  derived  from  a  previous  ameba;  it  mani- 
fests the  properties  and  performs  the  functions  of  its  life  which  have  been 
already  enumerated.  In  the  higher  organisms  it  is  different.  Each,  indeed, 
begins  as  a  single  cell,  but  the  cells  which  result  from  division  and  subdivision 
do  not  form  so  many  independent  organisms  but  adhere  in  one  differentiated 
community  which  ultimately  forms  the  complex  but  co-ordinated  whole,  in 
man  the  human  body. 

Thus  from  the  ovum  or  germ  cell  which  forms  the  starting-point  of  an 
individual  animal  during  development,  there  is  rapidly  formed  a  number  of 
tissues  each  characterized  by  its  own  type  of  structure,  the  whole  laid  down  in 
an  orderly  manner  to  form  the  complicated  individual.  In  the  unfolding 
of  this  individual  growth  process,  the  developing  ovum  soon  forms  a  complete 
membrane  of  cells  called  the  blastoderm,  and  this  speedily  differentiates  into 
two  and  then  into  three  layers,  chiefly  from  the  rapid  proliferation  of  the  cells 
of  the  first  single  layer.  These  layers,  figure  10,  are  called  the  Epiblast,  the 
Mesoblast,  and  the  Hypoblast.  In  the  further  development  of  the  animal 


DIFFERENCES    BETWEEN    ANIMALS    AND    PLANTS 


II 


it  is  found  that  from  each  of  these  layers  is  produced  a  very  definite  part 'of 
the  completed  body.  For  example,  from  the  cells  of  the  epiblast  are  derived, 
among  other  structures,  the  skin  and  the  central  nervous  system;  from  the 
mesoblast  the  muscles  and  connective  tissue  of  the  body,  and  from  the 
hypoblast  the  epithelium  of  the  alimentary  canal,  some  of  the  chief  glands, 
and  so  on. 

The  result  of  this  developmental  process  therefore  is  the  formation  of  the 
adult  tissues  highly  differentiated  and  specialized  in  form. 

From  the  physiologist's  point  of  view  this  anatomical  differentiation 
accomplishes  a  highly  specialized  structure  which,  machine-like,  is  capable  of 
doing  some  part  of  the  total  work  of  the  body  in  an  especially  effective  manner. 
In  a  word,  the  differentiated  tissues  do  not  altogether  lose  the  general  proper- 
ties which  characterize  protoplasm,  but  each  tissue  develops  a  structure  capa- 
ble of  doing  some  special  part  of  this  activity  better  than  the  undifferentiated 
protoplasm  can  do  it.  As  an  illustration,  the  muscles,  derived  chiefly  from 


FIG.  10. — Transverse  Section  through  Embryo  Chick  (26  hours),  a,  Epiblast;  b, 
mesoblast;  c,  hypoblast;  d,  central  portion  of  mesoblast,  which  is  here  fused  with  epiblast; 
e,  primitive  groove;  /,  dorsal  ridge.  (Klein.) 

either  epithelial  cells  or  mesoblast,  are  highly  contractile,  and  especially 
responsive  to  stimuli.  They  have  not  developed  in  the  same  degree  the  power 
to  produce  chemical  substances  which  characterizes  the  salivary  glands.  The 
cells  of  the  liver,  on  the  other  hand,  in  the  adult  stage  have  practically  lost  the 
property  of  contractility,  but  have  developed  in  a  high  degree  the  functional 
properties  of  nutrition  and  secretion. 

Hand  in  hand  with  the  anatomical  differentiation  has  gone  physiological 
division  of  labor.  In  the  adult  animal  body  each  type  of  activity  is  no  longer 
accomplished  by  the  whole  organism  as  in  the  case  of  the  ameba,  but  now 
some  specializing  part  of  the  body  assumes  each  chief  activity.  In  other 
words  the  muscles  contract,  the  nervous  tissue  conducts  changes  from  one  part 
of  the  body  to  the  other,  glands  secrete,  special  sense  organs  respond  to  the 
stimuli  of  the  environment,  and  the  reproductive  gonads  have  assumed  the 
chief  responsibility  in  the  reproductive  process.  It  is  through  the  high 
degree  of  physiological  division  of  labor  that  the  great  versatility  of  the  body 
activity  is  accomplished  by  the  human  organism.  It  is  a  matter  of  great 
economy  and  effectiveness  in  this  biological  machine,  the  body. 


12 


THE    PHENOMENA    OF    LIFE 


Differences  between  Animals  and  Plants. — Having  considered  the 
vital  properties  of  protoplasm,  as  shown  in  cells  of  animal  as  well  as  of  vege- 
table organisms,  we  are  now  in  a  position  to  discuss  the  question  of  the  differ- 
ences between  plants  and  animals.  It  might  at  the  outset  of  our  inquiry  have 
seemed  an  unnecessary  thing  to  recount  the  distinctions  which  exist  between 
an  animal  and  a  vegetable  organism  as  they  are  in  many  cases  so  obvious,  but, 
however  great  the  differences  may  be  between  the  higher  animals  and  plants, 
in  the  lowest  of  them  the  distinctions  are  much  less  plain. 

In  the  first  place,  it  is  important  to  lay  stress  upon  the  differences  between 
vegetable  and  animal  cells,  first  as  regards  their  structures  and  next  as  re- 
gards their  functions. 

It  has  been  already  mentioned  that  in  animal  cells  an  envelope  or  cell  wall 
is  by  no  means  always  present.  In  adult  vegetable  cells,  on  the  other  hand, 
a  well-defined  wall  is  highly  characteristic;  this  is  composed  of  cellulose, 
is  non-nitrogenous,  and  thus  differs  chemically  as  well  as  structurally  from 
the  contained  protoplasmic  mass.  Moreover,  in  vegetable  cells,  figure  n, 
B,  the  protoplasmic  contents  of  the  cell  fall  into  two  subdivisions:  i,  a  con- 
tinuous film  which  lines  the  interior  of  the  cellulose  wall;  and,  2,  a  reticulate 


FIG.  ii. — A.  Young  Vegetable  Cells,  Showing  Cell  Cavity  Entirely  Filled  with  Gran- 
ular Protoplasm  Enclosing  a  Large  Oval  Nucleus,  with  one  or  more  Nucleoli.  B.  Older 
cells  from  same  plant,  showing  distinct  cellulose  wall  and  vacuolation  of  protoplasm. 

mass  containing  the  nucleus  and  occupying  the  cell  cavity.  The  inter- 
stices are  filled  with  fluid.  In  young  vegetable  cells  such  a  distinction  does 
not  exist;  a  finely  granular  protoplasm  occupies  the  whole  cell  cavity,  figure 
n,  A.  As  regards  the  respective  functions  of  animal  and  vegetable  cells, 
one  of  the  most  important  differences  consists  in  the  power  which  vegetable 
cells  possess  of  being  able  to  build  up  new  complicated  nitrogenous  and 
non-nitrogenous  bodies  out  of  very  simple  chemical  substances  obtained 
from  the  air  and  from  the  soil.  They  obtain  from  the  air  oxygen,  carbon 
dioxide,  and  water,  as  well  as  traces  of  ammonia  gas;  and  from  the  soil  they 
obtain  water,  ammonium  salts,  nitrates,  sulphates,  and  phosphates  in  com- 
bination with  such  bases  as  potassium,  calcium,  magnesium,  sodium,  iron, 
and  others.  The  majority  of  plants  are  able  to  work  up  these  elementary 
compounds  into  other  and  more  complicated  bodies.  This  they  are  able 


DIFFERENCES    BETWEEN    ANIMALS    AND    PLANTS  13 

to  do  in  consequence  of  their  containing  a  certain  coloring  matter  called 
chlorophyll,  the  presence  of  which  is  the  cause  of  the  green  hue  of  plants. 
In  all  plants  which  contain  chlorophyll  two  processes  are  constantly  going 
on  when  they  are  exposed  to  light:  one,  which  is  called  true  respiration  and 
is  a  process  common  to  animal  and  vegetable  cells  alike,  consists  in  the 
taking  of  the  oxygen  from  the  atmosphere  and  the  giving  out  of  carbon 
dioxide;  the  other,  which  is  peculiar  apparently  to  bodies  containing  chloro- 
phyll, consists  in  the  taking  in  of  carbon  dioxide  and  the  giving  out  of  oxygen. 
It  seems  that  the  chlorophyll  is  capable  of  decomposing  the  carbon  dioxide 
gas  and  of  fixing  the  carbon  in  the  structures  in  the  form  of  new  compounds, 
one  of  the  most  rapidly  formed  of  which  is  starch. 

Vegetable  protoplasm  by  the  aid  of  its  chlorophyll  is  able  to  build  up  a 
large  number  of  bodies  besides  starch,  the  most  interesting  and  important 
being  protein  or  albumin.  It  appears  to  be  a  fact  that  the  power  which 
plants  possess  of  elaborate  chemical  synthesis  is  to  a  large  extent  dependent 
upon  the  chlorophyll  they  contain.  Thus  the  power  is  present  to  a  marked 
extent  only  in  the  plants  in  which  chlorophyll  is  found,  and  is  absent  in 
those  saphrophytic  plants  which  do  not  possess  chlorophyll. 

It  must  be  recollected,  however,  that  chlorophyll  without  the  aid  of  the 
light  of  the  sun  can  do  nothing  in  the  way  of  building  up  substances,  and  a 
plant  containing  chlorophyll  when  placed  in  the  dark,  while  it  continues  to 
live,  though  not  as  a  rule  long,  acts  as  though  it  did  not  contain  any  of  that 
substance.  It  is  an  interesting  fact  that  certain  of  the  bacteria  have  the 
chlorophyll  replaced  by  a  similar  pigment  which  is  able  to  decompose  carbon 
dioxide  gas. 

Animal  cells  do  not  possess  the  power  of  building  up  or  synthesizing  from 
simple  materials,  though  higher  organic  synthesis  can  no  longer  be  ques- 
tioned. Their  activity  is  chiefly  exercised  in  the  opposite  direction,  viz.,  the 
oxidations  of  the  complicated  compounds  produced  by  the  vegetable  kingdom 
which  they  have  brought  to  them  as  foods.  With  these  foods  animals  are  able 
to  perform  their  complex  functions,  setting  free  energy  in  the  direction  of 
heat,  motion,  and  electricity,  and  at  the  same  time  eliminating  such  bodies  as 
carbon  dioxide  and  water,  and  producing  other  bodies,  many  of  which  contain 
nitrogen  but  are  derived  from  decomposition. 

With  reference  to  the  substance  chlorophyll  it  has  been  noted  that  'the 
synthetical  operations  of  vegetable  cells  are  peculiarly  associated  with  the 
possession  of  chlorophyll  and  that  these  operations  are  dependent  upon  the 
light  of  the  sun.  It  has  been  further  shown  that  a  solution  of  chlorophyll 
when  examined  with  the  spectroscope  reveals  a  definite  absorption  spectrum, 
and  that  it  is  particularly  those  parts  of  the  solar  spectrum  corresponding 
to  these  absorption  bands  which  are  chiefly  active  in  the  decomposition  of 
carbon  dioxide.  In  the  synthetical  processes  of  the  plant,  then,  by  aid  of  its 
chlorophyll,  the  radiant  energy  of  the  sun's  rays  becomes  stored  up  or  ren- 


14  THE    PHENOMENA    OF   LIFE 

dered  potential  in  the  chemical  products  formed.  The  potential  energy  is 
set  free,  or  is  again  made  kinetic,  when  these  products  by  simple  combustion 
produce  heat,  or  when  they  are  taken  into  the  animal  organism  and  used  as 
food  and  there  later  produce  heat  and  motion. 

The  influence  of  light  is  not  absolutely  essential  to  animal  life;  indeed, 
it  is  said  not  to  increase  the  metabolism  of  animal  tissue  to  any  great  extent, 
and  the  animal  cell  does  not  receive  its  energy  directly  from  the  sun's  light 
nor  yet  to  any  extent  from  the  sun's  heat,  but  from  the  potential  energy  of  the 
food  stuffs.  But  it  must  be  always  kept  in  mind  that  anabolism  is  not  pecu- 
liar to  vegetable,  or  katabolism  to  animal  cells;  both  processes  go  on  in  each. 
Some  of  the  lowest  forms  of  vegetable  life,  e.g.,  the  bacteria,  will  live  only  in  a 
highly  albuminous  medium,  and  in  fact  seem  to  require  for  their  growth 
elements  of  food  stuffs  which  are  essential  to  animal  life.  In  their  metabo- 
lism, too,  they  very  closely  approximate  animal  cells,  not  only  requiring  an 
atmosphere  of  oxygen,  but  giving  out  carbon  dioxide  freely,  and  secreting 
and  excreting  many  very  complicated  nitrogenous  bodies,  as  well  as  forming 
protein,  carbohydrates,  and  fat,  requiring  heat  but  not  light  for  the  due  per- 
formance of  their  functions.  However,  certain  bacteria  grow  only  in  the 
absence  of  oxygen. 

There  is,  commonly,  a  difference  in  general  chemical  composition  be- 
tween vegetables  and  animals,  even  in  their  lowest  forms;  for  associated 
with  the  protoplasm  of  the  former  is  a  considerable  amount  of  cellulose,  a 
substance  closely  allied  to  starch  and  containing  carbon,  hydrogen,  and 
oxygen  only.  The  presence  of  starch  in  vegetable  cells  is  very  character- 
istic, though,  as  we  have  seen  above,  it  is  not  distinctive,  and  a  substance, 
glycogen,  similar  in  composition  to  starch,  is  very  common  in  the  organs  and 
tissues  of  animals. 

Inherent  power  of  movement  is  a  quality  which  we  so  commonly  consider 
an  essential  indication  of  animal  nature  that  it  is  difficult  at  first  to  conceive 
of  its  existence  in  any  other.  The  capability  of  simple  motion  is  now  known, 
however,  to  exist  in  so  many  vegetable  forms  that  it  can  no  longer  be  held 
as  an  essential  distinction  between  them  and  animals,  and  ceases  to  be  a  mark 
by  which  one  can  be  distinguished  from  the  other.  Thus  the  zoospores  of 
many  of  the  Cryptogams  exhibit  ciliary  or  ameboid  movements  of  a  like 
kind  to  those  seen  in  amebae;  and  even  among  the  higher  orders  of  plants, 
many,  e.g.,Dioncca  muscipula  (Venus's  fly-trap),  and  Mimosa  sensitiva  (Sensi- 
tive plant)  exhibit  such  motion,  either  at  regular  times  or  on  the  applica- 
tion of  external  irritation.  Were  this  fact  taken  by  itself,  it  might  lead  one  to 
regard  them  as  sensitive  organisms.  Inherent  power  of  movement,  then, 
although  especially  characteristic  of  animal  nature,  is,  when  taken  by  itself, 
no  proof  of  it. 

Sources  and  Utilization  of  Physiological  Material. — In  studying  the 
functions  of  the  human  body  it  is  necessary  first  of  all  to  know  of  what  it  is 


CELL    DIFFERENTIATION  1 5 

composed,  of  what  tissues  and  organs  it  is  made  up;  this  can  of  course  be 
ascertained  only  by  the  dissection  of  the  dead  body,  and  thus  it  comes  that 
Anatomy,  the  science  which  treats  of  the  structure  of  organized  bodies,  is 
closely  associated  with  physiology,  which  treats  of  the  functions  of  these 
structures.  So  close,  indeed,  is  the  association  that  Histology,  which  is 
especially  concerned  with  the  minute  or  microscopic  structure  of  the  tissues 
and  organs  of  the  body  and  which  is,  strictly  speaking,  a  department  of 
anatomy,  is  often  included  in  works  on  physiology.  There  is  much  to  be 
said  in  favor  of  such  an  arrangement,  since  it  is  impossible  to  consider  the 
changes  which  take  place  in  any  tissue  during  life,  apart  from  the  knowledge 
of  the  structure  of  the  tissues  themselves.  There  is  indeed  an  almost  insep- 
arable relation  between  the  structure  and  the  function  of  the  differentiated 
animal  body  in  which  the  one  is  made  the  means  to  a  knowledge  of  the  other 
as  an  end,  and  vice  versa,  according  to  the  aims  and  purposes  of  the  student. 

An  equally  important  essential  to  the  right  comprehension  of  the  changes 
which  take  place  in  the  living  organism  is  a  knowledge  of  the  chemical  com- 
position of  the  body.  Here,  however,  we  can  deal  directly  only  with  the 
composition  of  the  dead  body,  and  it  is  well  at  once  to  admit  that  there  may 
be  many  chemical  differences  between  living  and  non-living  tissues;  but  as  it 
is  impossible  to  ascertain  the  exact  chemical  composition  of  the  living  tissues, 
the  next  best  thing  which  can  be  done  is  to  find  out  as  much  as  possible  about 
the  composition  of  the  same  tissues  after  they  are  dead.  This  is  the  assist- 
ance which  the  science  of  Chemistry  can  afford  to  the  physiologist. 

Having  considered  the  structure  and  composition  of  the  body,  we'  are 
brought  face  to  face  with  physiology  proper,  and  have  to  investigate  the  vital 
changes  which  go  on  in  the  tissues,  the  various  actions  taking  place  as  long 
as  the  organism  is  at  work.  The  subject  includes  not  only  the  observation 
of  the  manifest  processes  which  are  continually  taking  place  in  the  healthy 
body,  but  the  conditions  under  which  these  are  brought  about,  the  laws 
which  govern  them  and  their  effects. 

It  may  be  well  to  mention  as  a  preliminary  that  the  physiological  informa- 
tion which  we  have  at  our  disposal  has  been  derived  from  many  sources,  the 
chief  of  which  are  as  follows:  i,  From  actual  observation  of  the  various 
phenomena  occurring  in  the  human  body  from  day  to  day,  and  from  hour  to 
hour,  as,  for  example,  the  estimation  of  the  amount  and  composition  of  the 
ingesta  and  egesta,  the  respiration,  the  beat  of  the  heart,  and  the  like;  2,  from 
observations  upon  other  animals,  the  bodies  of  which  we  are  taught  by  com- 
parative anatomy  approximate  the  human  body  in  structure,  and  may  be 
supposed  to  be  similar  in  function;  3,  from  observations  of  the  changes 
produced  by  experiment  upon  the  various  processes  in  such  animals,  or  in  the 
organs  and  tissues  of  animals;  4,  from  observations  of  the  changes  in  the 
working  of  the  human  body  produced  by  diseases;  and  5,  from  observations 
upon  the  gradual  changes  which  take  place  in  the  functions  of  organs  when 


1 6  THE   PHENOMENA    OF   LIFE 

watched  in  the  embryo  from  their  earliest  beginnings  to  their  completed 
development. 

The  physiologist,  in  order  to  utilize  the  sources  of  material,  must  be 
familiar  with  the  gross  structure  of  the  animals  or  parts  of  animals  which  he 
proposes  to  use  in  experimental  procedure.  So  simple  a  matter  as  the  deter- 
mination of  arterial  blood  pressure  involves  familiarity  with  extensive  ana- 
tomical structure.  Experimental  procedure  must  also  draw  on  the  field  of 
microscopic  structure  or  histology,  and  many  of  the  most  instructive  bodies  of 
physiological  knowledge  have  come  directly  from  the  utilization  of  the  facts 
of  comparative  anatomy  and  of  biology.  The  problems  in  animal  nutrition 
which  are  under  such  extensive  investigation  at  the  present  time  require  for 
their  solution  not  only  the  use  of  the  most  complex  methods  of  chemistry, 
both  analytic  and  synthetic,  but  also  the  principles  and  methods  of  physics. 
Indeed,  since  the  work  of  Helmholz,  the  interpretation  of  physiological  phe- 
nomena by  means  of  physical  methods  and  laws  has  contributed  more  than 
any  other  means  toward  the  prominent  scientific  position  of  physiology  at  the 
present  time.  In  a  word,  physiology  must  utilize  the  facts  of  anatomy,  his- 
tology, biology,  physics,  and  chemistry  to  interpret  the  phenomena  of  life. 


CHAPTER  II 

CELL  DIFFERENTIATION  AND  THE  TYPICAL  STRUCTURE  OF 
THE  ELEMENTARY  TISSUES 

In  the  preceding  discussion  a  general  view  of  the  type  of  cell  activity  and 
the  structural  basis  therefor  has  been  briefly  presented.  Emphasis  has  been 
laid  on  the  fact  that  the  complicated  phenomena  of  life  are  manifested  through 
the  agency  of  the  tissues  and  cells.  The  histological  cells,  alone  or  in  com- 
bination, are  capable  of  all  the  activities  manifested  by  the  living  body. 
Throughout  the  different  phases  of  the  physiological  discussions  which  follow 
it  will  be  assumed  that  the  reader  has  some  knowledge  of  this  structural  basis. 
However,  for  the  purpose  of  reference  there  is  presented  in  this  chapter  a 
brief  but  elementary  review  of  the  characteristic  cytological  structure  of  the 
tissues  and  cells  of  the  animal  body. 

THE  ESSENTIAL  STRUCTURE  OF  THE  TYPICAL  CELL 

The  typical  cell  is  a  spherical  or  ovoid  mass  of  protoplasm.  It  is  of 
microscopic  size  and  varies  from  6  or  7  micra  in  diameter  for  the  lymphocytes 
and  erythrocytes  to  150  to  200  micra  for  the  diameters  of  the  larger  cell 
bodies  of  the  neurones.  Its  structure  is  quite  complex,  but  the  most  general 
differentiation  is  into  the  cell-mass  or  cytoplasm,  and  its  contained  nucleus. 
The  cytoplasm  is  sometimes  bounded  by  a  definite  cell  membrane,  but  in 
differentiated  animal  tissues  this  membrane  is  usually  not  present. 

The  Cell  Body. — The  cell  body  or  ctyoplasm  is  a  complex  semi-fluid  mass, 
the  determination  of  the  detailed  relations  of  which  has  presented  many 
difficulties.  The  cell  cytoplasm  is  usually  described  as  having  a  framework 
of  spongioplasm  supporting  a  homogeneous  hyaloplasm.  In  some  cells  there 
are  formed  materials  resulting  from  the  cellular  activity  called  metaplasm, 
figure  12. 

Cell  protoplasm  includes  several  kinds  of  stainable  granules  and  fibrils, 
some  are  essential  constituents  while  others  are  formed  by  the  reactions  of  the 
protoplasm  and  are  in  a  sense  extraneous  material.  These  structural 
features  are  made  more  evident  by  their  selective  affinity  for  certain  staining 
reagents. 

The  exact  form  of  the  spongioplasm  or  reticulum  varies  greatly  in  different 
types  of  cells,  and  even  in  different  parts  of  the  same  cell.  Its  affinity  for 
stains  discloses  a  fine  network,  the  reticulum,  which  increases  in  amount 
and  also  in  constancy  in  the  type  of  arrangement  in  the  older  cells. 

17 


i8 


CELL    DIFFERENTIATION   AND    THE    ELEMENTARY    TISSUES 


The  hyaloplasm  is  more  fluid,  less  refractile,  and  stains  with  great  diffi- 
culty. It  fills  the  interspaces  of  the  spongioplasm.  In  this  material  may 
be  embedded  such  substances  as  the  metaplasts  mentioned  above. 

The  hyaloplasm  contains  in  solution  the  various  nutritive  constituents 
brought  to  the  cell  as  well  as  the  soluble  end  products  of  its  chemical  activity. 
Here,  too,  are  found  the  various  hormones,  oxidases  and  enzymes  which 
play  so  important  a  part  in  the  cellular  reactions  in  the  different  types  of  cells. 

Structure  of  the  Nucleus. — The  nucleus  when  in  a  condition  of  rest 
is  bounded  by  a  distinct  membrane,  the  nuclear  membrane,  possibly  derived 
from  the  spongioplasm  of  the  cell,  which  encloses  the  nuclear  contents,  nucleo- 
plasm  or  karyoplasm.  The  membrane  consists  of  an  inner,  or  chromatic, 
and  of  an  outer,  or  achromatic  layer,  so  called  from  their  reaction  to  stains. 
The  nucleoplasm  is  made  up  of  a  reticular  network,  or  chromoplasm,  whose 
interspaces  are  filled  by  the  karyolymph,  or  nuclear  matrix,  a  homogeneous 
substance  which  is  rich  in  proteins,  has  but  slight  affinity  for  stains,  and  is 
supposed  to  be  fluid  in  consistency. 


Cell  membrane.   — 


Metaplasmic   gran-  <--'* 
ules. 


Karyosome  or  net-   •'--'< 

knob. 

Hyaloplasm. 

Spongioplasm.    


Linin  network. 
Nucleoplasm. 


. Attraction  sphere. 

Centrosome. 

.,  Plastids. 

'-.C\  -  - \--^,-~- V- .-  -----  Chromatin . 

--:--«r--.?V --"->-> Nuclear  membrane. 

•-  Nucleolus. 
\----s-~~jf--.        __  Vacuole. 


FIG.  12.— Diagram  of  a  Typical  Cell.     (Bailey.) 

The  network  is  composed  of  linin  or  achromatin,  a  transparent  unstain- 
able  framework,  and  of  chromatin,  which  stains  deeply.  It  is  supported  by 
the  linin,  and  occurs  sometimes  in  the  form  of  granules,  but  usually  as  irreg- 
ular anastomosing  threads,  both  thicker  primary  fibers  and  thinner  connect- 
ing branches.  The  threads  often  form  thickened  nodes,  karyosomes  or 
false  nucleoli,  at  their  points  of  intersection.  It  is  now  quite  generally  be- 
lieved that  the  chromatin  occurs  as  short,  rod-like,  and  highly  refractive 
masses,  which  are  embedded  in  the  linin  in  a  regular  series. 

The  nucleoli,  or  plasmosomes,  are  spherical  bodies  which  stain  deeply,  and 
may  either  lie  free  in  the  nuclear  matrix  or  be  attached  to  the  threads  of 
the  network. 


CELL    MULTIPLICATION  IQ 

The  Centrosome  and  Attraction  Sphere. — In  addition  to  the  nucleus, 
a  minute  spherical  body  called  the  centrosome  is  believed  to  be  constantly 
present  in  animal  cells,  though  sometimes  too  small  to  be  demonstrated. 
The  centrosome  is  smaller  than  the  nucleus,  close  to  which  it  lies,  and  exerts  a 
peculiar  attraction  for  the  protoplasmic  filaments  and  granules  in  its  vicinity, 
so  that  it  is  surrounded  by  a  zone  of  fine  radiating  fibrils,  forming  the  attrac- 
tion sphere  or  archoplasm.  Some  authorities  assert  that  the  centrosome 
lies  within  the  nucleus  in  the  resting  state,  and  passes  into  the  cell  proper  only 
in  the  earlier  stages  of  cell  division.  The  attraction  sphere  is  most  distinctly 
seen  in  cells  about  to  divide.  It  plays  an  important  role  in  nuclear  division, 
but  it  is  doubted  if  it  gives  the  initial  impulse  to  the  process. 

Cell  Multiplication. — Cells  increase  in  number  by  a  process  known 
as  cell  division,  of  which  the  first  act  is  nuclear  division.  In  fact  the  nucleus 
is  the  center  of  control  of  the  cell  mass  in  the  process  of  division.  Cell  multi- 
plication takes  place  by  two  recognized  methods,  direct  or  amitosis,  in  which 
there  is  little  disturbance  of  the  nuclear  network,  and  indirect  or  mitosis,  in 
which  there  is  a  complex  series  of  nuclear  network  changes. 


a. 


FIG.  13. — Akinesis,  Amitosis,  or  Direct  Cell  Division.  A,  Constriction  of  nucleus;  B, 
division  of  nucleus  and  constriction  of  cell  body;  C,  daughter  nuclei  still  connected  by  a 
thread,  division  being  delayed;  D,  division  of  cell  body  nearly  complete.  (After  Arnold.) 

Direct  Cell  Division  or  Amitosis. — The  division  of  a  cell  is  preceded 
by  division  of  its  nucleus.  Direct  or  simple  division,  amitosis  or  akinesis,  see 
figure  13,  occurs  without  any  change  in  the  arrangement  of  the  intranuclear 
network.  A  constriction  develops  at  the  center  of  the  nucleus,  possibly  pre- 
ceded by  division  of  the  nucleoli,  and  gradually  divides  it  into  two  equal 
daughter  nuclei.  A  similar  constriction  of  the  protoplasm  of  the  cell  occurs 
between  the  daughter  nuclei  and  divides  it  into  two  parts. 

Indirect  Cell  Division  or  Mitosis. — Indirect  division,  mitosis,  orkaryo- 
kinesis  is  the  usual  method  of  cell  division,  and  consists  of  a  series  of  changes 
in  the  arrangement  of  the  intranuclear  network,  resulting  in  the  exact  division 


2O 


CELL   DIFFERENTIATION  AND   THE   ELEMENTARY   TISSUES 


of  the  chromatic  fibers  into  two  parts,  which  form  the  chromoplasm  of  the 
daughter  nuclei.  The  changes  follow  a  closely  similar  course  in  both  plant 
and  animal  cells. 

The    process    may   be   divided   into   the   following 
stages: 

Prophase. — The  resting  nucleus  becomes  somewhat 
enlarged,  and  the  centrosome  (according  to  those  who 
regard  it  as  lying  normally  within  the  nucleus)  migrates 
into  the  cell  protoplasm.  The  centrosome  then  divides 
into  two  daughter  centrosomes  which  lie  near  the  nucleus 
but  are  separated  by  a  considerable  interval.  Each  is 
surrounded  by  the  radiating  fibrils  of  the  attraction 
sphere,  and  some  of  these  fibrils  pass  continuously  from 
one  centrosome  to  the  other,  forming  the  achromatic 
spindle.  At  the  same  time  the  intranuclear  network  be- 
comes converted  into  a  fine  convoluted  coil,  the  spirem  or 
skein,  which  may  be  either  continuous  or  else  broken  up 
into  several  threads.  The  thread  or  threads  then 
shorten  and  become  thicker,  while  the  convolutions, 
which  have  become  less  numerous,  arrange  themselves  in  a  series  of 
connecting  loops,  forming  the  wreath.  The  nuclear  membrane  and  the 
nucleolus  disappear,  the  latter  passing  at  times  into  the  cell  protoplasm  and 
disintegrating.  The  wreath  then  breaks  up  into  V-shaped  segments,  the 
chromosomes,  of  which  each  species  of  animal  has  a  constant  and  character- 
istic number.  This  varies  in  the  different  animals,  but  is  sixteen  in  man. 

The  two  centrosomes  migrate  to  the  poles  of  the  nucleus,  while  the  achro- 
matic spindle  which  connects  them  occupies  the  long  axis  of  the  nucleus 


FIG.  14. — Leuco- 
cyte of  Salamander 
Larva,  Showing  At- 
traction Sphere. 
(After  Flemming.) 


FIG.  15. — Early  Stages  of  Karyokinesis.  A.  The  thicker  primary  fibers  remain  and 
the  achromatic  spindle  appears.  B.  The  thick  fibers  split  into  two  and  the  achromatic 
spindle  becomes  longitudinal.  (Waldeyer.) 

The  chromosomes,  becoming  much  shorter  and  thicker,  gather  around  the 
spindle  in  its  equatorial  plane,  with  their  angles  directed  toward  the  center, 
forming  the  aster  or  monaster. 

Metaphase. — The  actual  division  of  the  nucleus  is  begun  at  this  time  by  the 
splitting  of  each  chromosome  longitudinally  into  halves  which  lie  at  first  close 
together  so  that  each  seems  doubled.  Soon  afterward  the  fibrils  of  the 


CELL    MULTIPLICATION 


21 


achromatic  spindle  begin  to  contract,  and  thus  separate  the  halves  of  the 
chromosomes  in  such  a  way  that  one-half  of  each  is  turned  toward  one  pole, 
and  the  other  half  toward  the  other.  As  this  continues,  the  two  groups, 
which  are  equal  in  size,  draw  away  from  each  other  and  from  the  equator, 
each  group  being  formed  of  daughter  chromosomes. 


FIG.  1 6. — Monaster  Stage  of  Karyokinesis.     (Rabl.) 

Anaphase. — The  two  groups  (daughter  chromosomes)  now  gradually  ap- 
proach their  respective  poles,  or  centrosomes,  and  the  equator  becomes  free. 
On  reaching  the  pole,  each  group  gathers  in  a  form  which  is  similar  in  arrange- 
ment to  the  monaster  and  is  known  as  the  diaster.  During  this  time  the  cell 
body  becomes  slightly  constricted  by  a  circular  groove  at  its  equatorial  plane. 

Telophase. — Soon  afterward  the  fibrils  of  the  chromatic  spindle  which 
connect  the  two  groups  begin  to  grow  dim  and  finally  disappear.  The  daugh- 


FIG.  17. — Stages  of  Karyokinesis.  A.  Commencing  separation  of  the  split  chromo- 
somes. B.  The  separation  further  advanced.  C.  The  separated  chromosomes  passing 
along  the  fibers  of  the  achromatic  spindle.  (Rabl.) 

ter  chromosomes  assume  the  form  of  threads  twisted  in  a  coil  and  develop 
each  a  nuclear  membrane  and  a  nucleolus,  forming  a  daughter  nucleus. 
The  nuclei  enlarge  and  the  nuclear  threads  assume  the  appearance  of  the 
resting  state  of  the  nucleus.  Meanwhile,  the  constriction  about  the  body 
of  the  cell  cytoplasm  has  become  deeper  and  deeper  until  the  protoplasm  is 
divided  into  two  equal  parts,  or  daughter  cells,  each  with  its  daughter  nucleus, 
and  the  process  of  karyokinesis  is  completed. 


22 


CELL   DIFFERENTIATION  AND   THE   ELEMENTARY   TISSUES 


The  Cell  Types. — All  of  the  elementary  tissues  consist  of  cells  and  of 
their  altered  equivalents.  It  will  be  as  well  therefore  to  indicate  some  of  the 
differences  between  the  cells  of  the  body.  They  are  named  in  various  ways, 
according  to  their  shape,  origin,  and  functions. 


Line  of  division  — "^| 

of  cells. 

Antipole  of  daughter  ' 
nucleus. 


—  Remains  of  spindle. 


"'-->  Lighter  substance 
--'~""        of  nucleus. 

Cell  protoplasm. 
Hilus. 


FIG.  1 8. — Final  Stages  of  Karyokinesis.     In  the  lower  figure  the  changes  are  still  more 
advanced  than  in  the  upper.     (Waldeyer.) 

From  their  shape,  cells  are  described  as  spherical  or  spheroidal,  which  is 
the  typical  shape  of  the  free  cell;  this  may  be  altered  to  polyhedral  when  the 
pressure  on  a  mass  of  cells  in  all  directions  is  nearly  the  same;  of  this  the 
primitive  segmentation  cells  afford  an  example.  The  discoid  form  is  seen 


FIG.  19. — Karyokinesis,  Mitosis,  or  Indirect  Cell  Division  (diagrammatic).  A,  Cell 
with  resting  nucleus;  B,  wreath,  daughter  centrosomes  and  early  stage  of  achromatic 
spindle;  C,  chromosomes;  D,  monaster  stage,  achromatic  spindle  in  long  axis  of  nucleus, 
chromosomes  dividing;  E,  chromosomes  moving  toward  centrosomes;  F,  diaster  stage, 
chromosomes  at  poles  of  nucleus,  commencing  constriction  of  cell  body;  C,  daughter  nuclei 
beginning  return  to  resting  state;  H,  daughter  nuclei  showing  monaster  and  wreath;  7, 
complete  division  of  cell  body  into  daughter  cells  whose  nuclei  have  returned  to  the  resting 
state.  (After  Bohm  and  von  Davidoff.) 

in  blood  corpuscles,  and  the  scale-like  form  in  superficial  epithelial  cells. 
Some  cells  have  a  jagged  outline  and  are  then  called  prickle  cells.  Cells  of 
cylindrical,  conical,  or  prismatic  form  occur  in  various  places  in  the  body. 
Such  cells  may  taper  at  one  or  both  ends  into  fine  processes,  in  the  former  case 


THE    EPITHELIAL    TISSUES  23 

being  caudate,  in  the  latter  fusiform,  or  they  may  be  greatly  elongated  so  as  to 
become  fibers.  Cells  with  hair-like  processes,  or  cilia,  projecting  from  their 
free  surfaces,  are  a  special  variety.  The  cilia  vary  greatly  in  size,  and  may 
even  exceed  in  length  the  cell  itself.  Finally,  cells  may  be  branched  or  stellate 
with  long  outstanding  processes. 

From  theirfunction  cells  are  called  secreting,  protective,  sensitive,  contractile, 
and  the  like. 

From  their  origin  cells  are  called  epiblastic  and  mesoblastic  and  hypoblastic, 
ectodermic,  mesodermic,  and  endodermic. 

Modes  of  Cell  Connection. — Cells  are  connected  together  to  form  tissues 
in  various  ways.  They  are  connected  by  means  of  a  cementing  intercellular 
substance.  This  is  probably  always  present  as  a  transparent,  colorless,  viscid, 
albuminous  substance,  even  between  the  closely  apposed  cells  of  epithelium; 
while  in  the  case  of  cartilage  it  forms  the  main  bulk  of  the  tissue,  and  the  cells 
only  appear  as  embedded  in,  not  as  cemented  together  by,  the  intercellular 
substance.  This  intercellular  substance  may  be  either  homogeneous  or 
fibrillated.  In  many  cases,  e.g.,  the  cornea,  it  can  be  shown  to  contain  a 
number  of  irregular  branched  cavities,  which  communicate  with  each  other, 
and  in  which  branched  cells  lie.  Nutritive  fluids  can  find  their  way  through 
these  branching  spaces  into  the  very  remotest  parts  of  a  non-vascular  tissue. 
The  basement  membrane,  membrana  propria  must  be  mentioned  as  a  special 
variety  of  intercellular  substance  which  is  found  at  the  base  of  the  epithelial 
cells  in  most  mucous  membranes,  and  especially  as  an  investing  tunic  of 
gland  follicles  which  determines  their  shape. 

Cells  are  connected  by  anastomoses  of  their  processes.  This  is  the  usual 
way  in  which  stellate  cells,  e.g.,  of  the  cornea,  are  united.  The  individuality 
of  each  cell  is  thus  to  a  great  extent  lost  by  its  connection  with  its  neighbors 
to  form  a  reticulum.  As  an  example  of  a  network  so  produced  we  may  cite 
the  anastomosing  cells  of  the  reticular  tissue  of  lymphatic  glands. 

The  intercellular  substance  sometimes  forms  so  great  a  part  of  the  tissue 
as  to  overshadow  the  cells  proper.  Examples  of  this  type  of  structure  are 
found  in  the  matrix  of  cartilage,  the  fibers  of  connective  tissue,  bone,  etc. 

Decay  and  Death  of  Cells. — There  are  two  chief  ways  in  which  the 
comparatively  brief  existence  of  cells  is  brought  to  an  end,  i.e.,  by  mechanical 
abrasion  and  by  chemical  transformation. 

The  various  epithelia  furnish  abundant  examples  of  mechanical  abrasion. 
As  it  approaches  the  free  surface,  the  epidermal  cell  becomes  more  and  more 
flattened  and  scaly  in  form  and  more  horny  in  consistency,  till  at  length  it  is 
simply  rubbed  off.  Hence  we  find  free  epithelial  cells  in  the  mucus  of  the 
mouth,  in  the  intestine,  and  in  the  genito-urinary  tract,  as  well  as  on  the  sur- 
face of  the  outer  skin. 

In  the  case  of  chemical  transformation  the  cell  contents  undergo  a 
degeneration  which,  though  it  may  sometimes  be  pathological,  is  very  often 


24  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 

a  normal  process.  Thus  we  have  cells  by  fatty  metamorphosis  producing 
oil  globules  in  the  secretion  of  milk,  fatty  degeneration  of  the  muscular  fibers 
of  the  uterus  after  the  birth  of  the  fetus.  Calcareous  degeneration  is  common 
in  the  cells  of  many  cartilages. 

As  the  cells  approach  decay  and  death  their  normal  physiological  processes 
diminish  in  intensity  and  finally  cease.  This  occurs  early  in  the  transforma- 
tion and  function  is  lost  before  the  cell  form  is  destroyed  beyond  recognition. 

THE  STRUCTURE  OF  THE  ELEMENTARY  TISSUES. 

In  the  differentiation  of  the  protoplasm  of  the  body,  great  masses  of 
cells  are  formed  of  the  same  elemental  structure  and  typical  functional  prop- 
erties. These  are  the  elementary  tissues.  The  tissues  alone  or  in  combina- 
tion in  varying  proportions  constitute  the  organs  of  the  body.  These  ele- 
mentary tissues  are:  The  Epithelial,  The  Connective,  The  Muscular,  and  The 
Nervous  Tissues.  To  these  four  some  would  add  a  fifth,  looking  upon  the 
Blood  and  Lymph,  containing,  as  they  do,  formed  elements  in  a  fluid  men- 
struum, as  a  distinct  tissue. 

I.  THE  EPITHELIAL  TISSUES. 

Epithelium  is  a  tissue  composed  almost  wholly  of  cells,  with  a  very 
small  amount  of  intercellular  substance  which  glues  the  cells  together. 
In  general  it  includes  all  those  cellular  membranes  which  cover  either  an 
external  or  an  internal  free  surface,  together  with  the  cellular  portions  of  the 
glands  which  are  connected  with,  or  developed  from,  these  free  surfaces. 

Epithelium  clothes  (i)  the  whole  exterior  surface  of  the  body,  forming 
the  epidermis  with  its  appendages;  becoming  continuous  at  the  chief  orifices 
of  the  body — nose,  mouth,  anus,  and  urethra — with  (2)  the  epithelium  which 
lines  the  whole  length  of  the  respiratory,  alimentary,  and  genito-urinary 
tracts,  together  with  the  ducts  and  secretory  cells  of  their  various  glands. 
Epithelium  also  lines  the  cavities  of  (3)  the  brain  and  the  central  canal  of  the 
spinal  cord,  (4)  the  serous  and  synovial  membranes,  and  (5)  the  interior  of 
blood  vessels  and  lymphatics. 

Epithelial  cells  vary  in  size  and  shape,  pressure  being  the  main  factor  in 
this  variation.  The  protoplasm  may  be  granular,  reticular,  or  fibrillar  in 
appearance.  The  nucleus  is  spherical  or  oval,  usually  there  is  only  one,  but 
there  may  be  two  or  more  present. 

Epithelial  tissues  are  non-vascular,  that  is  to  say,  do  not  contain  blood 
vessels,  but  in  some  varieties  minute  channels  exist  between  the  cells  of 
certain  layers.  Nerve  fibers  are  supplied  to  the  cells  of  many  epithelia. 

CLASSIFICATION  OF  EPITHELIA. 
As  to  form  and  arrangement  of  cells. 

I.  Epithelia  in  the  form  of  membranes  (covering  surfaces), 
i.  Simple  epithelium.     Cells  only  one  layer  in  thickness. 


SIMPLE    EPITHELIUM  25 

(1)  Squamous  or  pavement.     Cells  flattened. 

(a)  Non-ciliated.     Alveoli  of  lungs,  also  includes  endothelium, 
lining  the  blood  vessels,  and  mesothelium,  lining  the  large 
serous  spaces. 

(b)  Ciliated.     The  peritoneum  of  some  forms  at  breeding  season. 

(2)  Cubical  epithelia.     Cells  with   the  three   dimensions   approxi- 

mately equal,  mainly  glandular. 

(a)  Non-ciliated.     The    usual  type.     It  is    found  lining    both 
ducts  and  secretory  portions  of  most  glands,  the  pigmented 
layer  of  the  retina,  etc. 

(b)  Ciliated.     Not  common.     Lining  of  some   of  the   smaller 
bronchial  tubes. 

(3)  Columnar.     Cells  may  be  cylindrical,  conical,  or  goblet-shaped. 

(a)  Non-ciliated.     Intestinal. 

(b)  Ciliated.     Fallopian  tube  and  uterus. 

(c)  Pseudo-stratified.     Smaller  bronchi,  nasal  duct,  etc. 
2.  Stratified  epithelia.     Cells  more  than  one  layer  in  thickness. 

(1)  Squamous.     Surface  cells  flattened. 

(a)  Non-ciliated.     Skin,  mouth,  vagina,  etc. 

(b)  Ciliated.     Pharynx  of  embryo. 

(2)  Columnar.     Surface  cells  columnar. 

(a)  Non-ciliated.     Portions  of  male  urethra. 

(b)  Ciliated.     Trachea,  bronchi,  etc. 

II.  Epithelia  not  in  the  form  of  membranes,  but  in  solid  masses  or  cords, 
usually  glandular. 

(1)  Cells  spheroidal.     Ova. 

(2)  Cells  polyhedral.     Liver,  suprarenal,  etc. 
Epithelia,  classified  mainly  as  to  function. 

I.  Protective.     Skin,  mouth,  alimentary  canal. 

1.  Cornified.     Skin,  nails,  hair. 

2.  Cuticular  border.     Columnar  cells  of  intestine. 
II.  Glandular. 

1.  Secretory.     Cells  of  salivary  glands,  pancreas,  etc. 

2.  Excretory.     Cells  of  kidney. 

3.  Absorptive.     Cells  of  alimentary  canal. 

III.  Sensory  Epithelium.     Cells  of  olfactory  membrane,  organ  of  Corti, 

taste  buds,  etc. 

IV.  Reproductive.     Sex  cells. 

V.  Pigmented.     Pigmented  layer  of  retina. 
VI.  Ciliated.     Trachea,  uterus,  Fallopian  tube,  etc. 
Only  a  few  of  the  more  important  of  the  above-mentioned  types  of  epithe- 
lium will  be  described  here. 

Simple    Epithelium. — Simple   Squamous. — This    form    of    epithelium 


20  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 

is  found  arranged  in  a  single  layer  of  flattened  cells,  for  example,  the  lining  of 
the  alveoli  of  the  lungs  and  of  the  descending  arm  of  Henle's  loop  of  the 
kidney  tubule.  Aside  from  endothelium  as  mesothelium  it  has  very  limited 
distribution  in  man.  Endothelium  and  mesothelium  are  typical  simple 
.squamous  epithelia.  They  consist  of  much  flattened  cells  with  clear  or 
.slightly  granular  protoplasm  and  oval  bulging  nuclei,  the  edges  of  the  cells 
.are  peculiarly  wavy  or  serrated. 


FIG.  20. — The  Endothelium  of  a  Small  Blood  Vessel.     Silver-nitrate  stain.      X  350. 

The  presence  of  endothelium  in  any  locality  may  be  demonstrated  by 
staining  with  silver  nitrate,  which  brings  into  view  the  intercellular  cement 
substance.  When  a  small  portion  of  a  perfectly  fresh  serous  membrane, 
for  example,  figure  20,  is  immersed  for  a  few  minutes  in  a  solution  of  silver 
nitrate,  and  exposed  to  the  action  of  light,  the  silver  is  precipitated  in  the  in- 
tercellular cement  substance,  and  the  endothelial  cells  are  thus  mapped  out 
by  fine,  dark,  and  generally  sinuous  lines  of  extreme  delicacy. 


FIG.  21. — Abdominal  Surface  of  Central  Tendon  of  the  Diaphragm  of  Rabbit,  showing  the 
general  polygonal  shape  of  the  endothelial  cells;  each  cell  is  nucleated.     (Klein.)      X  300. 

Endothelial  cells  in  certain  situations  may  be  ciliated,  e.g.,  those  of  the 
mesentery  of  the  frog,  especially  during  the  breeding  season. 

On  those  portions  of  the  peritoneum  and  other  serous  membranes  in 
which  lymphatics  abound,  apertures,  figure  22,  are  found  surrounded  by 
small,  more  or  less  cubical,  cells.  These  apertures  are  called  stomata.  They 
are  particularly  well  seen  in  the  anterior  wall  of  the  great  lymph  sac  of  the 
frog,  figure  22,  and  in  the  omentum  of  the  rabbit.  These  are  really  the  open 
mouths  of  lymphatic  vessels  or  spaces,  and  through  them  lymph  corpuscles 
and  the  serous  fluid  from  the  serous  cavity  pass  into  the  lymphatic  system. 


SIMPLE    EPITHELIUM  27 

Simple  Non-ciliated  Columnar  Epithelium,  figure  23,  lines,  a,  the  mucous 
membrane  of  the  stomach  and  intestines  as  a  single  layer,  from  the  cardiac 
orifice  of  the  stomach  to  the  anus,  and  b,  wholly  or  in  part  all  the  ducts  of  the 
glands  opening  on  its  free  surface,  and  c,  many  gland  ducts  in  other  regions 
of  the  body,  e.g.,  mammary,  salivary,  etc.  The  intracellular  and  intra- 


FiG.  22. — Peritoneal  Surface  of  a  Portion  of  the  Septum  of  the  great  Lymph  Sac  of  Frog. 
The  stomata,  some  of  which  are  open,  some  collapsed,  are  surrounded  by  endothelial  cells 
Klein.)  X  160. 

nuclear  networks  are  well  developed,  and  in  some  cases  the  spongioplasm  is 
arranged  in  rods  or  longitudinal  striae  at  one  part  of  the  cell,  as  in  the  cells  of 
the  ducts  of  salivary  glands.  The  protoplasm  of  columnar  cells  may  be 
vacuolated  and  may  also  contain  fat  or  other  substances  seen  in  the  form  of 
granules.  Certain  columnar  cells  transform  a  large  part  of  their  protoplasm 


FIG.  23. — Simple  Columnar  Epithelial  Cells  from  the  Human  Intestinal  Mucous 
Membrane,  a,  Mucous  (goblet)  cell;  b,  basement  membrane;  c,  cuticle;  d,  leucocyte 
nucleus;  e,  germinating  cell.  (Bailey.) 

into  mucin,  goblet  cells,  figure  24,  which  is  discharged  by  the  open  mouth 
of  the  goblet,  leaving  only  a  nucleus  surrounded  by  the  remains  of  the  proto- 
plasm in  its  narrow  stem.  This  transformation  is  a  normal  process  which 
is  continually  going  on  during  life,  the  cells  themselves  being  supposed  to 
regenerate  into  their  original  shape. 

Stratified  Epithelium. — The  term  stratified  epithelium  is  employed 


28 


CELL   DIFFERENTIATION  AND    THE   ELEMENTARY   TISSUES 


to  describe  the  type  found  in  the  skin  or  its  derivatives  in  which  the  cells 
forming  the  epithelium  are  arranged  in  a  considerable  number  of  superim- 
posed layers.  The  shape  and  size  of  the  cells  of  the  different  layers,  as  well 
as  the  number  of  layers,  vary  in  different  situations.  Thus  the  superficial 
cells  may  be  either  squamous  or  columnar  in  shape  and  the  deeper  cells 
range  from  polygonal  to  columnar  in  form. 


FIG.  24.  FIG.  25. 

FIG.  24.— Goblet  Cells.     (Klein.) 

FIG.  25. — Cross-section  of  a  Villus  of  the  Intestine,  e,  Columnar  epithelium  with 
striated  border;  g,  goblet  cell,  with  its  mucus  partly  extruded;  /,  lymph  corpuscles  between 
the  epithelial  cells;  &,  basement  membrane;  c,  sections  of  blood  capillaries;  m,  section  of 
plain  muscle  fibers;  c  I,  central  lacteal.  (Schafer.) 

Stratified  Squamous. — The  intermediate  cells  are  polygonal  in  shape  and 
approach  more  to  the  flat  variety  the  nearer  they  are  to  the  surface,  and  to  the 
columnar  as  they  approach  the  lowest  layer.  In  many  of  the  deeper  layers 
of  epithelium  in  the  mouth  and  skin,  the  outline  of  the  cells  is  very  irregular, 
in  consequence  of  processes  passing  from  cell  to  cell  across  these  intervals. 
Such  cells,  figure  28,  are  termed  "prickle"  cells.  These  "prickles"  are  the 


FlG.  26. — Squamous  Epithelium  Scales  from  the  Inside  of  the  Mouth.     X  260.     (Henle.) 

intercellular  bridges  which  run  across  from  cell  to  cell,  the  interstices  being 
filled  by  the  transparent  intercellular  lymph.  When  this  increases  in  quan- 
tity in  inflammation  the  cells  are  pushed  further  apart,  and  the  connecting 
fibrils  or  "prickles"  are  elongated  and  more  clearly  visible. 

The  columnar  cells  of  the  deepest  layer  are  distinctly  nucleated;  they 
multiply  rapidly  by  division;  and  as  new  cells  are  formed  beneath,  they  press 
the  older  cells  forward,  to  be  in  turn  pressed  forward  themselves  toward  the 


STRATIFIED    EPITHELIUM 


29 


surface,  gradually  altering   in   shape  and  chemical  composition  until  they 
die  and  are  cast  off  from  the  surface. 

Stratified  squamous  epithelium  is  found  in  the  following  situations:  i. 
Forming  the  epidermis,  covering  the  whole  of  the  external  surface  of  the  body; 
2.  Covering  the  mucous  membrane  of  the  nose,  tongue,  mouth,  pharynx,  and 
esophagus;  3.  As  the  conjunctival  epithelium,  covering  the  cornea;  4. 
Lining  the  vagina  and  the  vaginal  part  of  the  cervix  uteri. 


FIG.  27. — Vertical  Section  of  the  Stratified  Epithelium  Covering  the  Front  of  the 
Cornea.  Highly  magnified.  (Schafer.)  c,  Lowermost  columnar  cells;  p,  polygonal  cells 
above  these ;  fl,  flattened  cells  near  the  surface.  The  intercellular  channels,  bridged  by 
minute  cell  processes,  are  well  seen. 

Stratified  Columnar  Epithelium. — In  this  type  of  epithelium,  the  surface 
cells  alone  are  columnar,  the  deeper  cells  being  irregular  in  shape.  From 
the  surface  cells  long  processes  extend  down  among  the  underlying  cells. 
This  type  of  epithelium  is  usually  ciliated,  as  in  the  trachea,  bronchi,  etc., 
but  may  be  non-ciliated,  as  in  portions  of  the  human  male  urethra. 


a^i^^^'^^M 

FIG.  28. — Epithelial  Cells  from  the  Stratum  Spinosum  of  the  Human  Epidermis,  Showing 
"Intercellular  Bridges."     X  700.     (Szymonowicz.) 

Transitional  Epithelium. — This  is  a  stratified  epithelium  consisting  of 
only  three  or  four  layers  of  cells.  The  superficial  cells  are  large  and  flat, 
often  containing  two  nuclei.  The  under  surfaces  of  these  cells  are  hollowed 
out,  and  into  these  depressions  fit  the  large  ends  of  the  pyriform  cells  which 
form  the  next  layer.  Beneath  the  layer  of  pyriform  cells  are  from  one  to 


30  CELL    DIFFERENTIATION   AND    THE    ELEMENTARY    TISSUES 

four  layers  of  polyhedral  cells.     This  type  of  epithelium  occurs  in  the  bladder, 
ureter,  and  pelvis  of  the  kidney. 

Specialized  Epithelium. — Glandular  epithelium  forms  the  active  secreting 
agent  in  the  glands;  the  cells  are  usually  spheroidal,  but  may  be  polyhedral 
from  mutual  pressure,  or  even  columnar;  their  protoplasm  is  generally  oc- 
cupied by  the  materials  which  the  gland  secretes.  Examples,  of  glandular 


FIG.  29.  —  Stratified  Columnar  Ciliated  Epithelium  from  the  Human  Trachea. 

(goblet)  cell  also  is  present. 


A  mucous 


epithelium  are  to  be  found  in  the  liver,  figure  31,  in  the  secreting  tubes  of 
the  kidney,  and  in  the  salivary,  figure  32,  and  gastric  glands. 

Ciliated  epithelium  consists  of  cells  which  are  generally  cylindrical  in  form, 
figures  29,  30,  but  may  be  spheroidal  or  even  squamous. 

This  form  of  epithelium  lines:  a.  The  mucous  membrane  of  the  respira- 
tory tract  beginning  just  beyond  the  nasal  aperture,  and  completely  covers 
the  nasal  passages,  except  the  upper  part  to  which  the  olfactory  nerve  is 


. 

- 


FIG.  30.  —  Transitional  Epithelium  from  the  Human  Bladder.     (Bailey.) 

distributed,  and  also  the  sinuses  and  ducts  in  connection  with  it  and  the 
lachrymal  sac,  the  upper  surface  of  the  soft  palate  and  the  naso-pharynx, 
the  Eustachian  tube  and  tympanum,  the  larynx,  except  over  the  vocal  cords, 
to  the  finest  subdivisions  of  the  bronchi.  In  part  of  this  tract,  however, 
the  epithelium  is  in  several  layers,  of  which  only  the  most  superficial  is  ciliated, 


STRATIFIED    EPITHELIUM 


so  that  it  should  more  accurately  be  termed  transitional,  page  24,  or  stratified. 
b.  Some  portions  of  the  generative  apparatus  in  the  male,  viz.,  lining  the 
vasa  efferentia  of  the  testicle,  and  their  prolongations  as  far  as  the  lower 


FIG.  31. 


FIG.  32. 


FIG.  31. — A  Small  Piece  of  the  Liver  of  the  Horse.     (Cadiat.) 

FIG.  32. — Glandular  Epithelium.     Small  lobule  of  a  mucous  gland  of  the  tongue, 
showing  nucleated  glandular  cells.     X  200.     (V.  D.  Harris.) 

end  of  the  epididymis,  and  much  of  the  vas  deferens;  in  the  female,  c,  com- 
mencing about  the  middle  of  the  neck  of  the  uterus,  and  extending  throughout 
the  uterus  and  Fallopian  tubes  to  their  fimbriated  extremities,  and  even  for 


FIG.  33. — Specialized  Pigmented  Epithelial  Cells  of  Retina. 

a  short  distance  on  the  peritoneal  surface  of  the  latter,  d.  The  ventricles  of 
the  brain  and  the  central  canal  of  the  spinal  cord  are  clothed  with  ciliated 
epithelium  in  the  child,  but  in  the  adult  this  epithelium  is  limited  to  the  cen- 
tral canal  of  the  cord. 

The  cilia  themselves  are  fine  rounded  or  flattened  homogeneous  processes. 


32 


CELL    DIFFERENTIATION    AND    THE    ELEMENTARY    TISSUES 


According  to  some  observers,  these  processes  are  connected  with  longitudinal 
fibers  which  pass  to  the  other  end  of  the  cell,  but  which  are  not  connected 
with  the  nucleus. 


FIG.  34.  FIG.  35. 

FIG.  34. — Spheroidal  Ciliated  Cells  from  the  Mouth  of  the  Frog.  X  300  diameters. 
(Sharpey.) 

FIG.  35. — Ciliated  Epithelium  from  the  Human  Trachea,  a,  Large,  fully  formed  cell; 
ft,  shorter  cell;  c,  developing  cells  with  more  than  one  nucleus.  (Cadiat.) 

Functions  of  Epithelium. — According  to  function, 
epithelial  cells  may  be  classified  as:  i,  protective,  e.g.,  in 
the  skin,  mouth,  blood  vessels,  etc.;  2,  protective  and  motive, 
ciliated  epithelium;  3,  secreting,  glandular  epithelium;  4, 
germinal,  as  epithelium  of  testicle  producing  spermatozoa; 
5,  absorbing  and  secreting,  e.g.,  epithelium  of  intestine;  6, 
sensory,  e.g.,  olfactory  cells,  organ  of  Corti. 

Epithelium  forms  a  continuous  smooth  investment 
over  the  whole  body,  being  thickened  into  a  hard,  horny 
tissue  at  the  points  most  exposed  to  pressure,  and  develop- 
ing various  appendages,  such  as  hairs  and  nails.  Epithe- 
lium lines  also  the  sensorial  surfaces  of  the  eye,  ear,  nose, 
and  mouth,  and  thus  serves  as  the  medium  through  which 
all  impressions  from  the  external  world — touch,  smell, 
taste,  sight,  hearing — reach  the  delicate  nerve  endings, 
whence  they  are  conveyed  to  the  brain. 

The  ciliated  epithelium  which  lines  the  air  passages 
serves  to  promote  currents  of  the  air  in  the  bronchial  tubes 
and  to  propel  fluids  and  minute  particles  of  solid  matter 
out  of  the  body.  In  the  case  of  the  Fallopian  tube,  the 
cilia  assist  the  progress  of  the  ovum  toward  the  cavity  of 
the  uterus. 

The  epithelium  of  the  various  glands,  and  of  the 
(En-  whole  intestinal  tract,  has  the  power  of  secretion,  i.e.,  of 


FIG.  36. — Cili- 
ated Cell  of  the 
Intestine  of  a 
Mollusk. 

gelmann.) 

producing    certain 

ism  in  its  protoplasm. 


materials    by    processes    of    metabol- 


THE    CONNECTIVE    TISSUE  33 

Epithelium  is  likewise  concerned  in  the  processes  of  transudation, 
diffusion,  and  absorption. 

II.  THE  CONNECTIVE  TISSUES. 

This  group  of  tissues  forms  the  skeleton  with  its  various  connections — 
bones,  cartilages,  and  ligaments — and  also  affords  a  supporting  framework 
and  investment  to  the  various  organs  composed  of  nervous,  muscular,  and 
glandular  tissue.  Its  chief  function  is  the  mechanical  one  of  support,  and 
for  this  purpose  it  is  so  intimately  interwoven  with  nearly  all  the  textures  of 
the  body  that  if  all  other  tissues  could  be  removed,  and  the  connective  tissues 
left,  we  should  have  a  wonderfully  exact  model  of  almost  every  organ  and 
tissue  in  the  body. 

General  Structure  of  Connective  Tissue. — The  connective  tissue  is 
made  up  of  two  chief  elements,  namely,  cells  and  intercellular  or  formed 
substance. 


FIG.  37. — Horizontal  Preparation  of  the  Cornea  of  Frog,  Stained  in  Gold  Chloride; 
showing  the  network  of  branched  corneal  corpuscles.  The  ground  substance  is  completely 
colorless.  X  400.  (Klein.) 

Cells. — The  cells  are  usually  of  an  oval  shape,  often  with  branched 
processes,  which  are  united  to  form  a  network.  They  are  most  readily 
observed  in  the  cornea,  in  which  they  are  arranged,  layer  above  layer,  parallel 
to  the  free  surface.  They  lie  in  spaces  in  the  intercellular  or  ground  sub- 
stance, which  form  by  anastomosis  a  system  of  branching  canals  freely 
communicating,  figure  37. 

The  flattened  tendon  corpuscles  which  are  arranged  in  long  lines  or  rows 
parallel  to  the  fibers  belong  to  this  class  of  cells,  figure  39. 

These  branched  cells  often  contain  pigment  granules,  giving  them  a  dark 
appearance;  they  form  one  variety  of  pigment  cell.  Pigment  cells  of  this 
kind  are  found  in  the  outer  layers  of  the  choroid.  In  many  of  the  lower  ani- 
3 


34 


CELL    DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 


mals,  such  as  the  frog,  they  are  found  widely  distributed  not  only  in  the  skin, 
but  also  in  internal  parts,  the  mesentery,  sheaths  of  blood  vessels,  etc.  Under 
the  action  of  light,  electricity,  and  other  stimuli,  the  pigment  granules  become 
massed  in  the  body  of  the  cell,  leaving  the  processes  quite  hyaline;  if  the 
stimulus  be  removed,  they  will  gradually  be  distributed  again  throughout 
the  processes.  Thus  the  skin  in  the  frog  is  sometimes  uniformly  dusky  and 
sometimes  quite  light  colored,  with  isolated  dark  spots. 

Intercellular  Substance. — This  isjibrillar,  as  in  the  fibrous  tissues  and  in 
certain  varieties  of  cartilage;  or  homogeneous,  as  in  typical  mucoid  tissue. 

The  fibers  composing  the  former  are  of  two  kinds,  white  fibrous  and 
yellow  elastic  tissue. 


FIG.  38. 

FIG.  38. — Mature  White  Fibrous  Tissue  of  Tendon,  Consisting  Mainly  of  Fibers  with  a 
Few  Scattered  Fusiform  Cells.  (Strieker.) 

FIG.  39. — Caudal  Tendon  of  Young  Rat,  Showing  the  Arrangement,  Form,  and 
Structure  of  the  Tendon  Cells.  X  300.  (Klein.) 

The  chief  varieties  of  connective  tissues  may  be  thus  classified: 

White  fibrous. 

Elastic. 

Areolar. 

Gelatinous. 

Adenoid  or  retiform. 

Adipose. 

Neuroglia. 

Cartilage. 

1.  Hyaline. 

2.  White  fibrous. 

3.  Elastic. 
Bone  and  dentine. 

The  White  Fibrous  Tissue.— It  is  found  typically  in  tendon;  also  in 
ligaments,  in  the  periosteum  and  perichondrium,  the  dura  mater,  the  peri- 


GELATINOUS    TISSUE 


35 


cardium,  the  sclerotic  coat  of  the  eye,  the  fibrous  sheath  of  the  testicle,  in  the 
fasciae  and  aponeuroses  of  muscles,  and  in  the  sheaths  of  lymphatic  glands. 

Structure. — To  the  naked  eye,  tendons  and  many  of  the  fibrous  mem- 
branes, when  in  a  fresh  state,  present  an  appearance  as  of  watered  silk. 
This  is  due  to  the  arrangement  of  the  fibers  in  wavy  parallel  bundles.  Under 
the  microscope  the  tissue  appears  to  consist  of  long,  often  parallel,  bundles 
of  fibers  of  different  sizes.  The  cells  in  tendons,  figure  39,  are  arranged 
in  long  chains  in  the  ground  substance  separating  the  bundles  of  fibers,  and 
are  more  or  less  regularly  quadrilateral  with  large  round  nuclei  containing 
nucleoli,  generally  placed  so  as  to  be  contiguous  in  two  cells.  Each  of 
these  cells  consists  of  a  thick  body  from  which  processes  pass  in  various 
directions  into,  and  partially  fill  up  the  spaces  between,  the  bundles  of 
fibers.  The  rows  of  cells  are  separated  from  one  another  by  lines  of  cement 
substance. 

Yellow  Elastic  Tissue. — Yellow  elastic  tissue  is  found  chiefly  in  the 
ligamentum  nuchae  of  the  ox,  horse,  and  other  animals;  the  ligamenta  sub- 
flava  of  man;  the  arteries,  constituting  the  fenestrated  coat  of  Henle;  the 
veins  in  the  lungs  and  trachea;  the  stylo-hyoid,  thyro-hyoid,  and  crico- 
thyroid  ligaments;  in  the  true  vocal  cords;  and  in  areolar  tissue. 

Structure. — Elastic  tissue  occurs  in  various  forms,  from  a  structureless, 
elastic  membrane  to  a  tissue  whose  chief  constituents  are  bundles  of  fibers 
crossing  each  other  at  different  angles;  when  seen  in  bundles  elastic  fibers  are 
yellowish  in  color,  but  individual  fibers  are  not 
so  distinctly  colored.  The  varieties  of  the  tissue 
may  be  classified  as  follows: 

a.  Fine  elastic  fibrils,  which  branch  and  anas- 
tomose to  form  a  network.    This  variety  of  elastic 
tissue  occurs  chiefly  in   the  skin  and   mucous 
membranes,    in   subcutaneous    and    submucous 
tissue,  in  the  lungs  and  true  vocal  cords. 

b.  Thick  fibers,  sometimes  cylindrical,  some- 
times flattened,  which  branch,  anastomose  and 
form  a  network :  these  are  seen  most  typically  in 
the  ligamenta  subflava  and  also  in  the  ligamentum 
nuchae  of  such  animals  as  the  ox  and  horse,  in 
which  that  ligament  is  largely  developed,  figure  40. 

A  certain  number  of  connective-tissue  cells 
are  found  in  the  ground  substance  between 
the  elastic  fibers  which  make  up  this  variety  of 
connective  tissue,  page  33. 

Areolar  Tissue. — This  variety  of  fibrous  tissue  has  a  very  wide  dis- 
tribution and  constitutes  the  subcutaneous,  subserous,  and  submucous  tis- 
sue. It  is  found  in  the  mucous  membranes,  in  the  true  skin,  and  in  the  outer 


FIG.     40. — Elastic     Fibers 
from    the    Ligamenta    Sub- 
X  200.     (Sharpey.) 


CELL    DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 


sheaths  of  the  blood  vessels.  It  forms  sheaths  for  muscles,  nerves,  glands, 
and  the  internal  organs,  and,  penetrating  into  their  interior,  supports  and  con- 
nects the  finest  parts. 

Structure. — To  the  naked  eye  it  appears,  when  stretched  out,  as  a  fleecy, 
white,  and  soft  meshwork  of  fine  fibrils,  with  here  and  there  wider  films  join- 
ing in  it,  the  whole  tissue  being  evidently  elastic.  The  openness  of  the  mesh- 
work  varies  with  the  locality  from  which  the  specimen  is  taken.  Under  the 
microscope  it  is  found  to  be  made  up  of  fine  white  fibers,  which  interlace  in  a 
most  irregular  manner,  together  with  a  variable  number  of  elastic  fibers. 
On  the  addition  of  acetic  acid,  the  white  fibers  swell  up,  and  become  gelatin- 
ous in  appearance;  but  as  the  elastic  fibers  resist  the  action  of  the  acid,  they 
may  still  be  seen  arranged  in  various  directions,  sometimes  appearing  to  pass 
in  a  more  or  less  circular  or  spiral  manner  round  a  small  gelatinous  mass  of 
changed  white  fiber.  The  cells  of  areolar  tissues  are  connective-tissue 
corpuscles. 


V 

-4f 


FIG.  41.  FIG.  42. 

FIG.  41. — Mucous  Connective  Tissue  from  the  Umbilical  Cord,     a,  Cells;  b,  fibrils. 

FIG.  42. — Part  of  a  Section  of  a  Lymphatic  Gland,  from  which  the  corpuscles  have 
been  for  the  most  part  removed,  showing  the  Adenoid  Reticulum.  (Klein  and  Noble 
Smith.) 

Gelatinous  Tissue. — Gelatinous  connective  tissue  forms  the  chief 
part  of  the  bodies  of  such  marine  animals  as  the  jelly-fish.  It  is  found  in 
many  parts  of  the  human  embryo.  It  may  be  best  seen  in  the  "  Whartonian 
jelly"  of  the  umbilical  cord  and  in  the  enamel  organs  of  developing  teeth. 

Structure. — It  consists  of  cells,  which  in  the  jelly  of  the  enamel  organ 
are  stellate,  embedded  in  a  soft  jelly-like  intercellular  substance  which  forms 
the  bulk  of  the  tissue. 

Adenoid  or  Lymphoid  Tissue. — Distribution. — This  variety  of  tissue 
makes  up  the  stroma  of  the  spleen  and  lymphatic  glands,  and  is  found  also 


ADENOID    OR    LYMPHOID    TISSUE  37 

in  the  thymus,  in  the  tonsils,  and  in  the  follicular  glands  of  the  tongue;  in 
Peyer's  patches,  in  the  solitary  glands  of  the  intestines,  and  in  the  mucous 
membranes  generally. 

Structure. — Adenoid  or  retiform  tissue  consists  of  a  very  delicate  network 
of  minute  fibrils,  figure  42.  The  network  of  fibrils  is  concealed  by  being 
covered  with  flattened  connective-tissue  corpuscles,  which  may  be  readily 
dissolved  in  caustic  potash,  leaving  the  network  bare.  The  network  con- 
sists of  white  fibers,  the  interstices  of  which  are  filled  with  lymph  corpuscles. 
The  cement  substance  of  adenoid  tissue  is  very  fluid. 

Neuroglia. — This  form  of  connective  tissue  found  in  the  nervous  system 
is  described  on  page  78. 

Development  of  Fibrous  Tissues. — In  the  embryo  the  place  of  the  fibrous 
tissues  is  at  first  occupied  by  a  mass  of  roundish  cells,  derived 'chiefly  from 


FIG.  43. — Portion  of  Submucous  Tissue  of  Gravid  Uterus  of  Sow.     a,  Branched  cells, 
more  or  less  spindle-shaped;  b,  bundles  of  connective  tissue.     (Klein.) 

the  mesoderm,  but  also  from  ectoderm  and  from  entoderm.  These  develop 
either  into  a  network  of  branched  cells  or  into  groups  of  fusiform  cells, 
figure  43. 

The  cells  are  embedded  in  a  semifluid  albuminous  substance  derived 
probably  from  the  cells  themselves.  Later  this  formed  material  is  converted 
into  fibrils  under  the  influence  of  the  cells.  The  process  gives  rise  to  fibers 
arranged  in  the  one  case  in  interlacing  networks,  areolar  tissue,  in  the  other 
in  parallel  bundles,  white  fibrous  tissue.  In  the  mature  forms  of  purely 
fibrous  tissue  not  only  the  remnants  of  the  cell  substance,  but  even  the  nuclei, 
may  disappear.  The  embryonic  tissue,  from  which  elastic  fibers  are  devel- 
oped, is  composed  of  fusiform  cells  and  a  structureless  intercellular  sub- 
stance. The  fusiform  cells  dwindle  in  size  and  eventually  disappear  so 
completely  that  in  mature  elastic  tissue  hardly  a  trace  of  them  is  to  be  found; 
meanwhile  the  elastic  fibers  steadily  increase  in  size. 

Adipose  Tissue. — In  almost  all  regions  of  the  human  body  a  larger 
or  smaller  quantity  of  adipose  or  fatty  tissue  is  present.  Adipose  tissue  is 
almost  always  found  seated  in  areolar  tissue,  and  forms  in  its  meshes  little 
masses  of  unequal  size  and  irregular  shape,  to  which  the  term  lobules  is  com- 
monly applied. 


3&  CELL   DIFFERENTIATION   AND    THE    ELEMENTARY   TISSUES 

Structure. — Adipose  tissue  consists  essentially  of  cells  which  present 
dark,  sharply  denned  edges  when  viewed  with  transmitted  light;  each  con- 
sisting of  a  structureless  and  colorless  membrane  or  bag  formed  of  the  re- 
mains of  the  original  protoplasm  of  the  cell,  filled  with  fat.  A  nucleus 
is  always  present  in  some  part  or  other  of  the  cell  protoplasm,  but  in  the 
ordinary  condition  of  the  loaded  cell  it  is  not  easily  or  always  visible.  This 
membrane  and  the  nucleus  can  generally  be  brought  into  view  by  extracting 
the  fat  with  ether  and  by  staining  the  tissue. 


FIG.  44. — Blood  Vessels  of  Adipose  Tissue.  A,  Minute  flattened  fat  lobule,  in  which 
the  vessels  only  are  represented,  a,  The  terminal  artery;  v,  the  primitive  vein;  b,  the  fat 
vesicles  of  one  border  of  the  lobule  separately  represented.  X  100.  B,  Plan  of  the 
arrangement  of  the  capillaries,  c,  On  the  exterior  of  the  vesicles;  more  high  y  magnified. 
(Todd  and  Bowman.) 

The  ultimate  cells  are  held  together  by  capillary  blood  vessels,  figure  44; 
while  the  little  clusters  thus  formed  are  grouped  into  small  masses,  and 
held  so,  in  most  cases,  by  areolar  tissue.  The  oily  matter  contained  in  the 
cells  is  composed  chiefly  of  the  compounds  of  fatty  acids  with  gylcerin,  olein, 
stearin,  and  palmitin. 

Development  of  Adipose  Tissue. — Fat  cells  are  developed  from  connective- 
tissue  corpuscles.  In  the  infra-orbital  connective  tissue  there  are  cells  ex- 
hibiting every  intermediate  gradation  between  an  ordinary  branched  connect- 
ive-tissue corpuscle  and  mature  fat  cells.  Their  developmental  appearance 
is  as  follows:  a  few  small  drops  of  oil  make  their  appearance  in  the  proto- 
plasm, and  by  their  confluence  a  larger  drop  is  produced,  figure  45.  This 
gradually  increases  in  size  at  the  expense  of  the  original  protoplasm  of  the 
cell,  which  becomes  correspondingly  diminished  in  quantity  till  in  the  mature 
cell  it  forms  only  a  thin  crescentic  film  with  a  nucleus  closely  pressed  against 
the  cell  wall.  Under  certain  circumstances  this  process  may  be  reversed. 

A  large  number  of  blood  vessels  are  developed  in  adipose  tissue,  which 


CARTILAGE 


39 


subdivide  until  each  lobule  of  fat  contains  a  fine  meshwork  of  capillaries 
ensheathing  each  individual  fat  globule,  figure  44. 

Adipose  tissue  serves  as  a  storehouse  of  combustible  matter  which  may 
be  reabsorbed  into  the  blood  when  occasion  requires,  and,  being  used  up 
in  the  metabolism  of  the  tissues,  may  help  to  preserve  the  heat  of  the  body. 


FIG.  45. — A  Lobule  of  Developing 
Adipose  Tissue  from  an  Eight-months 
Fetus,  a,  Spherical  or,  from  pressure, 
polyhedral  cells  with  large  central  nu- 
cleus, surrounded  by  a  finely  reticulated 
substance  staining  uniformly  with  hema- 
toxylin.  b,  Similar  cells  with  spaces 
from  which  the  fat  has  been  removed 
by  oil  of  cloves,  c,  Similar  cells  showing 
how  the  nucleus  with  enclosing  proto- 
plasm is  being  pressed  toward  periphery. 
d,  Nucleus  of  endothelium  of  investing 
capillaries.  (McCarthy.) 


FIG.  46. — Branched  Connective- 
tissue  Corpuscles,  Developing 
into  Fat  Cells.  (Klein.) 


That  part  of  the  fat  which  is  situated  beneath  the  skin  must,  by  its  want  of 
conducting  power,  assist  in  preventing  undue  waste  of  the  heat  of  the  body 
by  escape  from  the  surface. 

CARTILAGE. 

All  kinds  of  cartilage  are  composed  of  cells  embedded  in  a  substance 
called  the  matrix.  The  apparent  differences  of  structure  met  with  in  the 
various  kinds  of  cartilage  are  more  due  to  differences  in  the  character  of 
the  matrix  than  of  the  cells.  With  the  exception  of  the  articular  variety, 
cartilage  is  invested  by  a  thin  but  tough  firm  fibrous  membrane  called  the 
perichondrium. 

Cartilage  exists  in  three  different  forms  in  the  human  body,  viz.,  hyaline 
cartilage,  yellow  elastic  cartilage,  and  white  fibro- cartilage. 

Hyaline  Cartilage. — This  variety  of  cartilage  is  met  with  largely  in 
the  human  body  where  it  invests  the  articular  ends  of  bones,  and  forms  the 


4O  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 

costal  cartilages,  the  nasal  cartilages,  and  those  of  the  larynx  with  the  ex- 
ception of  the  epiglottis  and  cornicula  laryngis,  the  cartilages  of  the  trachea 
and  bronchi. 


FIG.  47.  FIG.  48. 

FIG.  47. — Hyaline  Articular  Cartilage  (Human).     The  cell  bodies  entirely  fill  the  spaces 
in  the  matrix.      X  340  diams.     (Schafer.) 

FIG.  48. — Fresh  Cartilage  from  the  Triton.     (A.  Rollett.) 

Structure. — Like  other  cartilages,  it  is  composed  of  cells  embedded  in  a 
matrix.  The  cells  are  irregular  in  shape,  generally  grouped  together  in 
patches,  figure  47.  The  patches  are  of  various  shapes  and  sizes  and  placed 


FIG.  49. — Costal  Cartilage  from  an  Adult  Dog,  showing  the  Fat  Globules  in  the  Cartilage 

Cells.     (Cadiat.) 

at  unequal  distances  apart.  They  generally  appear  flattened  near  the  free 
surface  of  the  mass  of  cartilage,  and  more  or  less  perpendicular  to  the  surface 
in  the  more  deeply  seated  portions. 


ELASTIC    AND    WHITE    FIBRO-CARTILAGE  41 

The  intercellular  substance  of  hyaline  cartilage,  when  viewed  fresh  or 
after  ordinary  fixation,  appears  homogeneous.  However,  when  subjected 
to  special  methods,  the  seemingly  homogeneous  intercellular  substance  can 
be  shown  to  be  made  up  of  fibers,  comparable  with  those  found  in  white 
fibrous  tissue,  embedded  in  the  homogeneous  matrix. 

In  the  hyaline  cartilage  of  the  ribs  the  cells  are  mostly  larger  than  in 
the  articular  variety,  and  there  is  a  tendency  to  the  development  of  fibers 


ttliiil.1'1 


FIG.  50. — Yellow  Elastic  Cartilage  of  the 
Ear.     Highly  magnified.     (Hertwig.) 


FIG.  51. — White  Fibro-cartilage. 
(Cadiat.) 


in  the  matrix,  figure  49.  The  costal  cartilages  also  frequently  become 
calcified  in  old  age,  as  also  do  some  of  those  of  the  larynx. 

In  the  fetus  cartilage  is  the  material  of  which  the  bones  are  first  con- 
structed; the  "model"  of  each  bone  being  laid  down,  so  to  speak,  in  this 
substance.  In  such  cases  the  cartilage  is  termed  temporary.  It  closely 
resembles  the  ordinary  hyaline  cartilage,  but  the  cells  are  more  uniformly 
distributed  throughout  the  matrix. 

Elastic  and  White  Fibro-cartilage. — The  first  variety  is  found  in 
the  cartilage  of  the  external  ear;  the  latter  in  portions  of  the  joints,  the  inter- 
vertebral  cartilages,  etc. 

Structure. — Elastic  and  white  nbro-cartilage  are  composed  of  cells  and  a 
matrix;  the  latter  being  made  up  almost  entirely  of  fibers  closely  resembling 
those  of  fibrous  connective  tissue. 

Development  of  Cartilage. — Cartilage  is  developed  out  of  mesoblast  cells 
with  a  very  small  quantity  of  intercellular  substance.  The  cells  multiply  by 
fission  within  the  cell  capsules. 


42  CELL    DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 

BONE. 

The  characteristic  of  bone  is  that  the  matrix  is  solidified  by  a  deposit  of 
earthy  salts,  chiefly  calcium  phosphate,  but  some  magnesium  phosphate  and 
calcium  carbonate. 

To  the  naked  eye  there  appear  two  plans  of  structure  in  different  bones, 
and  in  different  parts  of  the  same  bone,  namely,  the  dense  or  compact,  and 
the  spongy  or  cancellous  tissue.  In  a  longitudinal  section  of  a  long  bone, 
as  the  humerus,  the  articular  extremities  are  found  capped  on  their  surface 
by  a  thin  shell  of  compact  bone,  while  their  interior  is  made  up  of  the  spongy 
or  cancellous  tissue.  The  shaft  is  formed  almost  entirely  of  a  thick  layer 
of  the  compact  bone  which  surrounds  a  central  canal,  the  medullary  cavity, 
so  called  from  its  containing  the  medulla,  or  marrow.  In  the  flat  bones,  the 
parietal  bone  or  the  scapula,  a  layer  of  cancellous  structure  lies  between 
two  layers  of  the  compact  tissue.  In  the  short  and  irregular  bones,  as  those 
of  the  carpus  and  tarsus,  the  cancellous  tissue  alone  fills  the  interior,  while 
a  thin  shell  of  compact  bone  forms  the  outside. 


FIG.  52. — Cells  of  the  Red  Marrow  of  the  Guinea-pig,  highly  magnified,  a,  A  large 
cell,  the  nucleus  of  which  appears  to  be  partly  divided  into  three  by  constrictions;  b,  a  cell, 
the  nucleus  of  which  shows  an  appearance  of  being  constricted  into  a  number  of  smaller 
nuclei;  c,  a  so-called  giant  cell,  or  myeloplaxe,  with  many  nuclei;  d,  a  smaller  myeloplaxe, 
with  three  nuclei;  e-i,  proper  cells  of  the  marrow.  (Schafer.) 

The  Marrow. — There  are  two  distinct  varieties  of  marrow — the  red  and 
the  yellow. 

Red  marrow  is  that  variety  which  occupies  the  spaces  in  the  cancellous 
tissue;  it  is  highly  vascular,  and  thus  maintains  the  nutrition  of  the  spongy 
bone,  the  interstices  of  which  it  fills.  It  contains  a  few  fat  cells  and  a  large 
number  of  marrow  cells,  many  of  which  are  undistinguishable  from 
lymphoid  corpuscles,  and  has  for  a  basis  a  small  amount  of  fibrous  tissue. 
Among  the  cells  are  some  nucleated  cells  containing  hemoglobin  like  the 
blood  corpuscles.  There  are  also  a  few  large  cells  with  many  nuclei,  termed 
giant  cells  or  myeloplaxes,  which  are  probably  derived  from  the  ordinary 
marrow  cells,  figure  52. 


THE    PERIOSTEUM    AND    NUTRIENT    BLOOD  VESSELS 


43 


Yellow  marrow  fills  the  medullary  cavity  of  long  bones,  and  consists 
chiefly  of  fat  cells  with  numerous  blood  vessels.  Many  of  its  cells  are  in 
every  respect  similar  to  lymphoid  corpuscles. 

From  these  marrow  cells,  especially  those  of  the  red  marrow,  the  red 
blood  corpuscles  are  derived. 

The  Periosteum  and  Nutrient  Blood  Vessels. — The  surfaces  of 
bones,  except  the  part  covered  with  articular  cartilage,  are  clothed  by  a 


FIG.  53. — Transverse  Section  of  Compact  Bone  (of  humerus).  Three  of  the  Haver- 
sian canals  are  seen,  with  their  concentric  rings;  also  the  lacunae,  with  the  canaliculi  extending 
from  them  across  the  direction  of  the  lamella.  •  The  Haversian  apertures  were  filled  with 
debris  in  grinding  down  the  section,  and  therefore  appear  black  in  the  figure,  which 
represents  the  object  as  viewed  with  transmitted  light.  The  Haversian  systems  are  so 
closely  packed  in  this  section,  that  scarcely  any  -interstitial  lamellce  are  visible.  X  150. 
(Sharpey.) 


tough,  fibrous  membrane,  the  periosteum,  which  is  closely  attached  to  the 
surface  of  the  bone.  Blood  vessels  are  distributed  in  this  membrane,  and 
minute  branches  from  these  periosteal  vessels  enter  the  Haversian  canals 
to  supply  blood  to  the  solid  part  of  the  bone.  The  long  bones  are  supplied 
also  by  a  proper  nutrient  artery  which,  entering  at  some  part  of  the  shaft 
so  as  to  reach  the  medullary  canal,  breaks  up  into  branches  for  the  supply 
of  the  marrow,  from  which  again  small  vessels  are  distributed  to  the  interior 
of  the  bone.  Other  small  nutrient  vessels  pierce  the  articular  extremities 
for  the  supply  of  the  cancellous  tissue. 


44  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 

Microscopic  Structure  of  Bone. — Notwithstanding  the  differences 
of  arrangement  just  mentioned,  the  structure  of  all  compact  bone  substance 
is  found  under  the  microscope  to  be  essentially  the  same. 

Examined  with  a  rather  high  power  its  substance  is  found  to  contain  a 
multitude  of  small  irregular  spaces,  approximately  fusiform  in  shape,  called 
lacuna,  with  very  minute  canals  or  canaliculi,  as  they  are  termed,  leading 


FIG.    54. — Longitudinal   Section  from   the   Human   Ulna,    Showing   Haversian   Canals, 
Lacunae,  and  Canaliculi.     (Rollett.) 


from  them,  and  anastomosing  with  similar  prolongations  from  other  lacunae, 
figure  53.  In  very  thin  layers  of  bone,  no  other  canals  than  these  may  be  vis- 
ible; but  on  making  a  transverse  section  of  the  compact  tissue  of  a  long  bone, 
as  the  humerus  or  ulna,  the  arrangement  shown  in  figure  53  can  be  seen. 
The  bone  seems  mapped  out  into  small  circular  districts,  at  or  about  the 
center  of  each  of  which  is  a  hole,  around  which  are  concentric  layers,  the 
lamella,  the  lacuna  and  canaliculi  following  the  same  concentric  distribution 
around  the  center,  with  which  indeed  they  communicate. 

On  making  a  longitudinal  section,  the  central  holes  are  shown  to  be 
simply  the  cut  extremities  of  small  canals  which  run  lengthwise  through 
the  bone,  anastomosing  with  each  other  by  lateral  branches,  figure  54,  and 
are  called  Haversian  Canals,  after  the  name  of  the  physician,  Clopton  Havers, 
who  first  accurately  described  them. 

The  Haversian  Canals. — The  average  diameter  of  the  Haversian  canals 
is  50^.  They  contain  blood  vessels,  and  by  means  of  them  blood  is  con- 
veyed to  even  the  densest  parts  of  the  bone;  the  minute  canaliculi  and  lacunae 


MICROSCOPIC    STRUCTURE    OF    BONE  45 

absorbing  nutrient  matter  from  the  Haversian  blood  vessels  and  conveying 
it  still  more  intimately  to  the  very  substance  of  the  bone  which  they  traverse. 
The  blood  vessels  enter  the  Haversian  canals  both  from  without  from  the 
periosteum,  and  from  within  from  the  medullary  cavity  or  from  the  can- 
cellous  tissue.  The  arteries  and  veins  usually  occupy  separate  canals. 

The  lacuncc  are  occupied  by  branched  cells,  the  bone  cells  or  bone  corpus- 
cles, figure  55,  which  very  closely  resemble  the  ordinary  branched  connective- 
tissue  corpuscles.  The  processes  of  the  bone  cells  extend  into  the  canaliculi. 
Each  cell  controls  the  nutrition  of  the  bone  immediately  surrounding  it. 
Each  lacunar  corpuscle  communicates  with  the  others  in  its  surrounding 


FIG.  55. — Bone  Corpuscles  with  their  Processes  as  seen  in  a  Thin  Section  of  Human  Bone. 

(Rollett.) 

district,  and  with  the  blood  vessels  of  the  Haversian  canals  by  means  of  the 
ramifications  just  described. 

It  will  be  seen  from  the  above  description  that  bone  bears  a  very  close 
structural  resemblance  to  what  may  be  termed  typical  connective  tissue. 
The  bone  corpuscles  with  their  processes  occupying  the  lacunae  and  canalic- 
uli correspond  exactly  to  the  cornea  corpuscles  lying  in  the  branched  spaces. 

The  Lamella  of  Compact  Bone. — In  the  shaft  of  a  long  bone  three  distinct 
sets  of  lamellae  can  be  clearly  recognized:  General  or  fundamental  lamellae, 
which  are  just  beneath  the  periosteum  and  parallel  with  it,  and  around  the 
medullary  cavity;  Special  or  Haversian  lamellae,  which  are  concentrically 
arranged  around  the  Haversian  canals  to  the  number  of  six  to  eighteen 
around  each;  Interstitial  lamellae,  which  connect  the  systems  of  Haversian 
lamellae,  filling  the  spaces  between  them,  and  consequently  attaining  their 
greatest  development  where  the  Haversian  systems  are  few. 

The  ultimate  structure  of  the  lamellae  appears  to  be  fibrous.  A  thin 
film  peeled  off  the  surface  of  a  bone,  from  which  the  earthy  matter  has  been 
removed  by  acid,  is  composed  of  a  finely  reticular  structure,  formed  ap- 


46  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 

parently  of  very  slender  fibers  decussating  obliquely,  but  coalescing  at  the 
points  of  intersection,  as  if  here  the  fibers  were  fused  rather  than  woven 
together.  The  reticular  lamellae  are  perforated  by  the  perforating  fibers  of 
Sharpey,  which  bolt  the  neighboring  lamellae  together,  and  may  be  drawn 
out  when  the  latter  are  torn  asunder,  figure  56.  These  perforating  fibers 
originate  from  ingrowing  processes  of  the  periosteum,  and  in  the  adult  still 
retain  their  connection  with  it. 


FIG.  56. — Lamellae  Torn  off  from  a  Decalcified  Human  Parietal  Bone  at  some  Depth 
from  the  Surface,  a,  a,  Lamellae,  showing  reticular  fibers;  b,  b,  darker  part,  where  several 
lamellae  are  superposed;  c,  perforating  fibers.  Apertures  through  which  perforating  fibers 
had  passed,  are  seen  especially  in  the  lower  part,  a,  a.  of  the  figure.  (Allen  Thomson.) 


Development  of  Bone. — From  the  point  of  view  of  their  development 
all  bones  may  be  subdivided  into  two  classes: 

Those  which  are  ossified  directly  in  membrane  or  fibrous  tissue,  e.g.,  the 
bones  forming  the  vault  of  the  skull,  parietal,  frontal,  and  a  certain  portion 
of  the  occipital  bones. 

Those  whose  form,  previous  to  ossification,  is  laid  down  in  hyaline  carti- 
lage, e.g.,  humerus,  femur,  etc. 

The  process  of  development,  pure  and  simple,  may  be  best  studied  in 
bones  which  are  not  preceded  by  cartilage,  i.e.,  membrane-formed.  Without 
a  knowledge  of  ossification  in  membrane  it  is  difficult  to  understand  the  much 
more  complex  series  of  changes  through  which  such  a  structure  as  the  carti- 
laginous femur  of  the  fetus  passes  in  its  transformation  into  the  bony  femur 
of  the  adult  (ossification  in  cartilage}. 

Ossification  in  Membrane. — The  membrane,  afterward  forming  the 
periosteum,  from  which  such  a  bone  as  the  parietal  is  developed,  consists 
of  two  layers,  an  external  fibrous  and  an  internal  cellular  or  osteogenetic. 


OSSIFICATION    IN    CARTILAGE  47 

The  external  layer  consists  of  ordinary  connective  tissue,  with  branched 
corpuscles  here  and  there  between  the  bundles  of  fibers.  The  internal  layer 
consists  of  a  network  of  fine  fibrils  with  nucleated  cells  and  ground  or  cement 
substance  between  the  fibrous  bundles.  It  is  more  richly  supplied  with 
capillaries  than  the  outer  layer.  The  relatively  large  number  of  its  cellular 
elements,  together  with  the  abundance  of  blood  vessels,  clearly  mark  it  as 
the  portion  of  the  periosteum  which  is  immediately  concerned  in  the  for- 
mation of  bone. 

In  such  a  bone  as  the  parietal  there  is  first  an  increase  in  vascularity, 
followed  by  the  deposition  of  bony  matter  in  radiating  spicula,  starting 
from  a  center  of  ossification.  These  primary  bony  spicula  are  osteo genetic 
•fibers,  composed  of  osteogen,  in  which  calcareous  granules  are  deposited. 
Calcareous  granules  are  deposited  also  in  the  interfibrillar  matrix.  By 
the  junction  of  the  osteogenetic  fibers  and  their  resulting  bony  spicula  a 
meshwork  of  bone  is  formed.  The  osteoblasts,  being  in  part  retained  within 
the  bone  trabeciilae  thus  produced,  form  bone  corpuscles.  Lime  salts  are 
deposited  in  the  circumferential  part  of  each  osteoblast,  and  thus  a  ring 
of  osteoblasts  gives  rise  to  a  ring  of  bone  with  the  remaining  uncalcified 
portions  of  the  osteoblasts  embedded  in  it  as  bone  corpuscles.  At  the  same 
time  the  plate  increases  at  the  periphery  by  the  extension  of  the  bony  spicula 
and  by  deposits  taking  place  from  the  osteogenetic  layer  of  the  periosteum. 
The  bulk  of  the  primitive  spongy  bone  is  gradually  converted  into  compact 
bony  tissue  of  the  Haversian  systems. 

Ossification  in  Cartilage. — Under  this  heading,  taking  the  femur  as 
a  typical  example,  we  may  consider  the  process  by  which  the  solid  cartilag- 
inous rod  which  represents  the  bone  in  the  fetus  is  converted  into  the  hollow 
cylinder  of  compact  bone  with  expanded  ends  formed  of  cancellous  tissue 
in  the  adult  long  bone. 

The  fetal  cartilage  is  sheathed  in  a  membrane  termed  the  perichondrium, 
which  resembles  the  periosteum  described  above.  Thus,  the  differences 
between  the  fetal  perichondrium  and  the  periosteum  of  the  adult  are  such 
as  usually  exists  between  the  embryonic  and  mature  forms  of  connective 
tissue. 

There  are  several  steps  in  the  transformation  of  the  fetal  cartilage  to  the 
adult  bone,  due  to  the  fact  that  there  is  first  an  impregnation  of  the  cartilage 
with  lime  salts,  followed  later  by  the  resorption  of  this  entire  material  with 
formation  of  the  embryonic  spongy  bone,  which  is  still  later  replaced  by  the 
permanent  bone.  The  complicated  phenomenon  takes  place  in  steps  or 
sagest  as  follows: 

Stage  of  Proliferation  and  Calcification. — The  cartilage  cells  in  and  near 
the  center  of  ossification  become  enlarged,  proliferate,  and  arrange  them- 
selves in  rows  in  the  long  axis  of  the  fetal  cartilage,  figure  57.  Lime  salts  are 
next  deposited  in  fine  granules  in  the  hyaline  matrix  of  the  cartilage,  and  this 


48 


CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 


gradually  becomes  transformed  into  calcified  trabeculse,  figure  57.  The  en- 
larging cartilage  cells  become  more  transparent,  and  finally  disintegrate,  the 
spaces  occupied  by  them  forming  the  primordial  marrow  cavities.  During 
this  stage  the  perichondrium  has  become  the  periosteum,  and  is  beginning 
to  deposit  bone  on  the  outside  of  the  cartilage. 


FIG.  57. — Developing  Bone  of  Femur  of  the  Rabbit.  X  350.  a,  Cartilage  cells; 
b,  cartilage  cells  enlarged  in  the  region  of  calcifying  matrix;  c,  d,  trabecuhc  of  calcifying 
cartilage  covered  with  e,  osteoblasts;  /,  osteocla'sts  eroding  the  trabecuke;  g,  h,  disap- 
pearing cartilage  cells.  The  osteoblasts  are  seen  to  be  depositing  layers  of  bony  sub- 
stance. Loops  of  blood  vessels  extend  to  the  limit  of  the  region  in  which  the  bone  is 
forming.  (Schafer,  from  Klein.) 

Stage  of  Vascular ization  of  the  Cartilage. — Processes  from  the  osteo- 
genetic  layer  of  the  periosteum  containing  blood  vessels  break  through  the 
bone  into  the  primordial  marrow  cavities  and  form  the  primary  marrow, 
beginning  at  the  centers  of  ossification,  and  spreading  chiefly  up  and  down 
the  shaft. 

Stage  of  Substitution  of  Embryonic  Spongy  Bone  for  Calcified  Cartilage. — 
The  cells  of  the  primary  marrow  arrange  themselves  as  a  continuous  epi- 


OSSIFICATION   IN    CARTILAGE 


49 


thelium-like  layer  on  the  calcined  trabeculae  and  deposit  a  layer  of  bone, 
and  ensheath  them.  The  encased  trabeculae  are  gradually  absorbed  by 
the  osteodasts  of  Kolliker. 

These  stages  are  precisely  similar  to  what  goes  on  in  the  growing  shaft 
of  a  bone  which  is  increasing  in  length  by  the  advance  of  the  process  of  ossifi- 


FiG.  58. — Transverse  Section  through  the  Tibia  of  a  Fetal  Kitten,  semidiagrammatic. 
X  60.  P,  Periosteum.  O,  Osteogenetic  layer  of  the  periosteum,  showing  the  osteoblasts 
arranged  side  by  side,  represented  as  pear-s'haped  black  dots  on  the  surface  of  the  newly 
formed  bone.  B,  The  periosteal  bone  deposited  in  successive  layers  beneath  the  peri- 
osteum and  ensheathing  E,  the  spongy  endochondral  bone;  represented  as  more  deeply 
shaded.  Within  the  trabeculae  of  endochondral  spongy  bone  are  seen  the  remains  of  the 
calcined  cartilage  trabeculae  represented  as  dark  wavy  lines.  C,  The  medulla,  with  V,  V, 
veins.  In  the  lower  half  of  the  figure  the  endochondral  spongy  bone  has  been  completely 
absorbed.  (Klein  and  Noble  Smith.) 

cation  into  the  intermediary  cartilage  between  the  diaphysis  and  epiphysis. 
In  this  case  the  cartilage  cells  become  flattened  and,  multiplying  by  division, 
are  grouped  into  regular  columns  at  right  angles  to  the  plane  of  calcification 
while  the  process  of  calcification  extends  into  the  hyaline  matrix  between 
them. 

4 


5° 


CELL    DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 


The  embryonic  spongy  bone,  formed  as  above  described,  is  simply  a  tem- 
porary tissue  occupying  the  place  of  the  fetal  rod  of  cartilage;  the  preceding 
stages  show  the  successive  changes  at  the  center  of  the  shaft.  Periosteal 
bone  is  at  the  same  time  deposited  in  successive  layers  beneath  the  perios- 
teum at  the  circumference  of  the  shaft,  exactly  as  described  in  the  section 
on  ossification  in  membrane,  and  thus  a  casing  of  periosteal  bone  is  formed 
around  the  embryonic  endochondral  spongy  bone.  The  embryonic  spongy 
bone  is  absorbed,  through  the  agency  of  the  osteoclasts,  until  the  trabeculas 
are  replaced  by  one  great  cavity,  the  medullary  cavity  of  the  shaft. 


FIG.  59. — Transverse  Section  of  Femur  of  a  Human  Embryo  about  Eleven  Weeks  Old. 
a,  Rudimentary  Haversian  canal  in  cross-section;  6,  in.  longitudinal  section;  c,  osteoblasts; 
d,  newly  formed  osseous  substance  of  a  lighter  color;  e,  that  of  greater  age;/,  lacunae  with 
their  cells;  g,  a  cell  still  united  to  an  osteoblast.  (Frey.) 


Stage  of  Formation  of  Compact  Bone. — The  transformation  of  spongy 
periosteal  bone  into  compact  bone  is  effected  in  a  manner  exactly  similar 
to  that  which  has  been  described  in  connection  with  ossification  in  mem- 
brane, page  46.  The  irregularities  in  the  walls  of  the  spongy  periosteal 
bone  are  absorbed  by  the  osteoclasts,  while  the  osteoblasts  which  line  them 
are  developed  in  concentric  layers,  each  layer  in  turn  becoming  ossified 
till  the  comparatively  large  space  in  the  center  is  reduced  to  a  well-formed 
Haversian  canal,  figure  59.  When  once  formed,  bony  tissue  grows  to  some 
extent  inter stitially,  as  is  evidenced  by  the  fact  that  the  lacunas  are  rather 
further  apart  in  full-formed  than  in  young  bone. 


THE    TEETH  51 

It  will  be  seen  that  the  common  terms  ossification  in  cartilage  and  ossifi- 
cation in  membrane  are  apt  to  mislead,  since  they  seem  to  imply  two  processes 
radically  distinct.  The  process  of  ossification,  however,  is  in  all  cases  one 
and  the  same,  all  true  bony  tissue  being  formed  from  membrane,  perichon- 
drium  or  periosteum;  but  in  the  development  of  such  a  bone  as  the  femur, 
lime  salts  are  first  of  all  deposited  in  the  cartilage;  this  calcified  cartilage, 
however,  is  gradually  and  entirely  reabsorbed,  replaced  by  bone  formed 
from  the  periosteum.  Thus  calcification  of  the  cartilaginous  matrix  pre- 
cedes the  real  formation  of  bone.  We  must,  therefore,  clearly  distinguish 
between  calcification  and  ossification.  The  former  is  simply  the  infiltration 
of  an  animal  tissue  with  lime  salts,  while  ossification  is  the  formation  of 
true  bone. 

Growth  of  Bone. — Bones  increase  in  length  by  the  advance  of  the 
process  of  ossification  into  the  cartilage  intermediate  between  the  diaphysis 
and  epiphysis.  The  increase  in  length  indeed  is  due  entirely  to  growth 
at  the  two  ends  of  the  shaft.  Increase  in  thickness  in  the  shaft  of  a  long 
bone  occurs  by  the  deposition  of  successive  layers  beneath  the  periosteum. 
If  a  thin  metal  plate  be  inserted  beneath  the  periosteum  of  a  growing  bone 
it  will  soon  be  covered  by  osseous  deposit,  but  if  it  be  put  between  the  fibrous 
and  osteogenetic  layers  it  will  never  become  enveloped  in  bone,  for  all  the 
bone  is  formed  beneath  the  latter. 


THE  TEETH. 

During  the  course  of  his  life,  man,  in  common  with  most  other  mammals, 
is  provided  with  two  sets  of  teeth;  the  first  set,  called  the  temporary  or  milk- 
teeth  of  infancy,  are  shed  and  replaced  by  the  second  or  permanent  set. 


Temporary  Teeth. 

MIDDLE  LINE  OF  JAW. 


Molars.        Canine.        Incisors. 

212 


Incisors.        Canine.        Molars. 

212         =IO 
212         =10 


The  figures  indicate  in  months  the  age  at  which  each  tooth  appears: 


Lower  central 
incisors 

Upper  incisors 

First  molar  and 
lower  lateral 
incisors 

Canines 

Second  molar 

6  to  9 

8  to  12 

I  2  to  I  5 

1  8  to  24 

24  to  30 

52 


CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 

Permanent  Teeth. 

MIDDLE  LINE  OF  JAW. 

Incisor,         Canine.       P^rT' 


The  age  at  which  each  permanent  tooth  is  cut  is  indicated  in  this  table  in  years 


First  molars 

Incisors 

Bicuspids  or 
premolars 

Canines 

Second 
molars 

Third 
molars  or 
wisdoms 

Centrals 

Laterals 

First 

Second 

6 

7 

8 

9 

IO 

12  to  14 

I  2  to  I  5 

17  to  2  5 

Structure. — A  tooth  is  generally  described  as  possessing  a  crown,  neck, 
and  root  or  roots.  The  crown  is  the  portion  which  projects  beyond  the 
level  of  the  gum.  The  neck  is  that  constricted  portion  just  below  the  crown 
which  is  embraced  by  the  free  edges  of  the  gum,  and  the  root  includes  all 
below  this. 

On  making  longitudinal  and  transverse  sections  through  its  center,  figure 
6 1,  A,  B,  a  tooth  is  found  to  be  principally  composed  of  a  hard  superficial 


FIG.  60. — Normal  Well-formed  Jaws,  from  which  the  Alveolar  Plate  has  been  in  great 
part  removed,  so  as  to  expose  the  Developing  Permanent  Teeth  in  their  Crypts  in  the  Jaws. 
(Tomes.) 

material,  dentine  or  ivory,  which  is  hollowed  out  into  a  central  cavity  which 
resembles  in  general  shape  the  outline  of  the  tooth,  and  is  called  the  pulp 
cavity. 

The  tooth  pulp  is  composed  of  fibrous  connective  tissue,  blood  vessels, 
nerves,  and  large  numbers  of  cells  of  varying  shapes.  On  the  surface  in 


DENTINE    OR    IVORY 


53 


close  connection  with  the  dentine  there  is  a  specialized  layer  of  cells  called 
odontoblasts,  which  are  elongated  columnar  cells  with  a  large  nucleus  at  the 
tapering  ends  farthest  from  the  dentine.  The  cells  are  all  embedded  in  a 
mucoid  gelatinous  matrix. 

The  blood  vessels  and  nerves  enter  the  pulp  through  a  small  opening 
at  the  apical  extremity  of  each  root. 

A  layer  of  very  hard  calcareous  matter,  the  enamel,  caps  the  dentine  of 
the  crown;  beneath  the  level  of  the  gum  is  a  layer  of  true  bone,  called  the 
cement  or  crusta  petrosa.  The  enamel  and  cement  are  very  thin  at  the  neck 
of  the  tooth  where  they  come  in  contact,  the  cement  overlapping  the  enamel. 
The  enamel  becomes  thicker  toward  the  crown,  and  the  cement  toward 
the  lower  end  or  apex  of  the  root. 


FIG.  61. — A.  A  Longitudinal  Section  of  a  Human  Molar  Tooth,  c,  Cement;  d, 
dentine;  e,  enamel;  v,  pulp  cavity.  B. — Transverse  section.  The  letters  indicate  the 
same  as  in  A  (Owen). 


Dentine  or  Ivory. — Dentine  closely  resembles  bone  in  chemical  com- 
position. It  contains,  however,  rather  less  animal  matter. 

Structure. — Dentine  is  finely  channelled  by  a  multitude  of  delicate  tubes, 
which  by  their  inner  ends  communicate  with  the  pulp  cavity,  and  by  their 
outer  extremities  come  into  contact  with  the  under  part  of  the  enamel  and 
cement,  and  sometimes  even  penetrate  them  for  a  greater  or  less  distance, 
figures  63,  64.  The  matrix  in  which  these  tubes  lie  is  composed  of  "a  retic- 
ulum  of  fine  fibers  of  connective  tissue  modified  by  calcification,  and, 
where  that  process  is  complete,  entirely  hidden  by  the  densely  deposited 
lime  salts"  (Mummery). 

The  tubules  of  the  dentine  contain  fine  prolongations  from  the  tooth 
pulp,  which  gives  the  dentine  a  certain  faint  sensitiveness  under  ordinary 
circumstances  and,  without  doubt,  have  to  do  also  with  its  nutrition.  They 
are  probably  processes  of  the  dentine  cells  or  odontoblasts  lining  the  pulp 


54 


CELL    DIFFERENTIATION   AND    THE    ELEMENTARY    TISSUES 


cavity.     The  relation  of  these  processes  to  the  tubules  in  which  they  lie  is 
precisely  similar  to  that  of  the  processes  of  the  bone  corpuscles  to  the  canalic- 


Dentine.    — 


Enamel. 


Cement. 


Periosteum  of 
alveolus. 


Cement. J 


Lower  jaw  bone. 


FIG.  62. — Premolar  Tooth  and  Surrounding  Bone  of  Cat. 

ali  of  bone.     The  outer  portion  of  the  dentine,  underlying  the  cement  and 
the  enamel,  figure  63,  6,  c,  contains  cells  like  bone  corpuscles. 


FIG.  63. — Section  of  a  Portion  of  the  Dentine  and  Cement  from  the  Middle  of  the  Root 
of  an  Incisor  Tooth,  a,  Dental  tubuli  ramifying  and  terminating,  some  of  them  in  the 
interglobular  spaces  b  and  c,  which  somewhat  resemble  bone  lacunse;  d,  inner  layer  of  the 
cement  with  numerous  closely  set  canaliculi;  e,  outer  layer  of  cement;/,  lacunae;  g,  canalic- 
uli.  X  350.  (Kolliker.) 

Enamel. — The  enamel,  which  is  by  far  the  hardest  portion  of  a  tooth, 
is  composed  chemically  of  the  same  elements  that  enter  into  the  composition 


ENAMEL 


55 


of  dentine  and  bone,  but  the  animal  matter  amounts  only  to  about  2  or  3 
per  cent.  It  contains  a  larger  proportion  of  inorganic  matter  and  is  harder 
than  any  other  tissue  in  the  body. 

Structure. — Enamel  is  composed  of  fine  hexagonal  fibers,  figures  64,  65. 


FIG.  64. 

FIG.  64. — Thin  Section  of  the  Enamel  and  a  Part  of  the  Dentine,  a,  Cuticular  pellicle 
of  the  enamel  (Nasmyth's  membrane);  b,  enamel  fibers,  or  columns  with  fissures  between 
them  and  cross  striae;  c,  larger  cavities  in  the  enamel,  communicating  with  the  extremities 
of  some  of  the  dentinal  tubuli  (d).  X  350.  (Kolliker.) 

FIG.  65. — Section  of  the  Upper  Jaw  of  a  Fetal  Sheep.  A. — i,  Common  enamel  germ 
dipping  down  into  the  mucous  membrane;  2,  palatine  process  of  jaw;  3,  rete  Malpighi. 
(Waldeyer.)  B. — Section  similar  to  A,  but  passing  through  one  of  the  special  enamel  germs 
here  becoming  flask-shaped;  c,  c,  epithelium  of  mouth;/,  neck;/',  body  of  special  enamel 
germ.  (Rose.)  C. — A  later  stage;  c,  outline  of  epithelium  of  gum;  /,  neck  of  enamel 
germ;  /',  enamel  organ;  p.  papilla;  s,  dental  sac  forming;//?,  the  enamel,  germ  of  perma- 
nent tooth;  m,  bone  of  jaw;  v,  vessels  cut  across.  (Kolliker.)  Copied  from  Quain's 
"Anatomy." 

These  are  set  on  end  vertical  to  the  surface  of  the  dentine,  and  fit  into  cor- 
responding depressions  in  the  same. 

Like  the  dentine  tubules,  they  are  disposed  in  wavy  and  parallel  curves. 


56  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 

The  fibers  are  thus  marked  by  transverse  lines.  They  are  mostly  solid, 
but  some  of  them  may  contain  a  very  minute  canal. 

The  enamel  prisms  are  connected  together  by  a  trace  of  hyaline  cement 
substance. 

Development. — The  first  step  in  the  development  of  the  teeth  consists 
in  a  downward  growth,  figure  65,  A,  i,  from  the  deeper  layer  of  stratified 
epithelium  of  the  mouth,  which  first  becomes  thickened  in  the  neighborhood 
of  the  maxillae  or  jaws,  now  also  in  the  course  of  formation.  This  epidermal 
papilla  grows  downward  into  a  recess  of  the  imperfectly  developed  tissue  of 
the  embryonic  jaw.  It  forms  the  primary  enamel  organ  or  enamel  germ,  and 
its  position  is  indicated  by  a  slight  groove  in  the  mucous  membrane  of  the 
jaw.  The  next  step  consists  in  the  elongation  and  the  inclination  outward 


FIG.  66. — Part  of  Section  of  Developing  Tooth  of  a  Young  Rat,  showing  the  Mode 
of  Deposition  of  the  Dentine.  Highly  magnified,  a,  Outer  layer  of  fully  formed  dentine; 

b,  uncalcified  matrix  with  one  or  two  nodules  of  calcareous  matter  near  the  calcined  parts; 

c,  odontoblasts  sending  processes  into  the  dentine;  d,  pulp;  e,  fusiform  or  wedge-shape  cells 
found  between  odontoblasts;  /,  stellate  cells  of  pulp  in  fibrous  connective  tissue.     The 
section  is  stained  in  carmine,  which  colors  the  uncalcified  matrix  but  not  the  calcified  part. 
(E.  A.  Schafer.) 

of  the  deeper  part,  figure  65,  B,  /',  of  the  enamel  germ,  followed  by  an  in- 
creased development  at  certain  points  corresponding  to  the  situations  of  the 
future  milk-teeth.  The  enamel  germ  becomes  divided  at  its  deeper  portion, 
or  extended  by  further  growth,  into  a  number  of  special  enamel  germs  corre- 
sponding to  each  of  the  milk-teeth,  and  connected  to  the  common  germ  by  a 
narrow  neck.  Each  tooth  is  thus  placed  in  its  own  special  recess  in  the 
embryonic  jaw,  figure  65,  c,  /'. 

As  these  changes  proceed,  tnere  grows  up  from  the  underlying  tissue 
into  each  enamel  germ,  figure  65,  c,  p,  a  distinct  vascular  papilla,  dental 
papilla,  and  upon  it  the  enamel  germ  becomes  molded,  and  presents  the 
appearance  of  a  cap  of  two  layers  of  epithelium  separated  by  an  interval, 
figure  65,  c,  /'.  While  part  of  the  subepithelial  tissue  is  elevated  to  form 
the  dental  papillae,  the  part  which  bounds  the  embryonic  teeth  forms  the 
dental  sacs,  figure  65,  c,  s;  and  the  rudiment  of  the  jaw  sends  up  processes 
forming  partitions  between  the  teeth.  The  papilla,  which  is  really  part  of 
the  dental  sac,  is  composed  of  nucleated  cells  arranged  in  a  meshwork.  in 


ENAMEL 


57 


the  outer  layer  of  which  are  the  columnar  cells  called  odontoblasts.  The 
odontoblasts  form  the  dentine,  while  the  remainder  of  the  papilla  forms  the 
pulp.  The  method  of  the  formation  of  the  dentine  from  the  odontoblasts 
is  said  to  be  as  follows:  The  cells  form  elongated  processes  at  their  outer 
surfaces  which  are  directly  converted  into  the  tubules  of  dentine,  figure  66,  c, 
and  into  the  contained  fibrils. 

Each  papilla  early  takes  the  shape  of  the  crown  of  the  tooth  to  which 
it  corresponds,  but  as  the  dentine  increases  in  thickness  and  papilla  dimin- 
ishes until  when  the  tooth  is  cut  only  a  small  amount  remains  as  the  pulp.  It 
is  supplied  by  vessels  and  nerves  which  enter  at  the  end  of  the  root.  The 
roots  are  not  completely  formed  at  the  time  of  the  eruption  of  the  teeth. 


FIG.  67.— Vertical  Transverse  Section  of  the  Dental  Sac,  Pulp,  etc.,  of  a  Kitten,  a, 
Dental  papilla  or  pulp;  b,  the  cap  of  dentine  formed  upon  the  summit;  c,  its  covering  of 
enamel;  d,  inner  layer  of  epithelium  of  the  enamel  organ;  e,  gelatinous  tissue;/,  outer 
epithelial  layer  of  the  enamel  organ;  g,  inner  layer,  and  h,  outer  layer  of  dental  sac.  X  14. 
(Thiersch.) 

The  enamel  cap  is  formed  by  the  enamel  cells,  by  the  deposit  of  a  keratin- 
like  substance,  which  subsequently  undergoes  calcification.  Other  layers 
are  formed  in  the  same  manner  meanwhile. 

The  temporary  or  milk-teeth  are  speedily  replaced  by  the  growth  of  the 
permanent  teeth. 

The  development  of  the  temporary  teeth  commences  about  the  sixth 
week  of  intra-uterine  life,  after  the  laying  down  of  the  bony  structure  of 
the  jaws.  Their  permanent  successors  begin  to  form  about  the  sixteenth 
week  of  intra-uterine  life. 


58  CELL    DIFFERENTIATION   AND    THE    ELEMENTARY   TISSUES 

III.  MUSCULAR  TISSUE. 

There  are  two  chief  kinds  of  muscular  tissue,  differing  both  in  minute 
structure  as  well  as  in  mode  of  action,  viz.,  (i)  the  smooth  or  non- striated,  and 
(2)  the  striated. 

SMOOTH  OR  NON-STRIATED  MUSCLE. 

Non-striated  muscle  forms  the  proper  muscular  coats  of  the  digestive 
canal  from  the  middle  of  the  esophagus  to  the  internal  sphincter  ani;  of 
the  uterus  and  urinary  bladder;  of  the  trachea  and  bronchi;  of  the  ducts 


FIG.  68. — Isolated  Smooth  Muscle  Cells  from  Human  Small  Intestine.  X  400.  Rod- 
shaped  nucleus  surrounded  by  area  of  finely  granular  protoplasm;  longitudinal  striations 
of  cytoplasm. 

of  glands;  of  the  gall-bladder;  of  the  vesiculae  seminales;  of  the  uterus  and 
Fallopian  tubes;  of  the  blood  vessels  and  lymphatics;  and  of  the  iris  and 
some  other  parts  of  the  eye.  This  form  of  tissue  also  enters  largely  into  the 
composition  of  the  tunica  dartos  of  the  scrotum.  Unstriped  muscular  tissue 


FIG.  69. — Smooth  Muscle  from  Intestine  of  Pig,  Showing  Syncytial  Structure,  a, 
Protoplasmic  process  connecting  two  muscle  fibers;  b,  end-to-end  union  of  two  muscle 
fibers,  showing  the  continuity  of  protoplasm  and  myofibrils;  c,  nucleus  of  muscle  fiber; 
d,  granular  protoplasm  at  the  end  of  muscle  nucleus;  e,  coarse  myofibril;/,  fine  myofibril; 
g,  connective-tissue  cell  with  connective-tissue  fibrils  surrounding  it;  h,  elastic  fiber.  (New 
figure  by  Caroline  McGill.) 

occurs  largely  also  in  the  true  skin  generally,  being  especially  abundant  in  the 
interspaces  between  the  bases  of  the  papillae,  and,  when  it  contracts,  the 
papillae  are  made  unusually  prominent,  giving  rise  to  the  peculiar  roughness 
of  the  skin  termed  cutis  anserina,  or  goose  flesh.  It  also  occurs  in  all  parts 


STRUCTURE 


59 


where  hairs  occur,  in  the  form  of  flattened  roundish  bundles  which  lie  along- 
side the  hair  follicles  and  sebaceous  glands. 

Structure. — Unstriated   muscle   fibers   are   elongated,   spindle-shaped 


FIG.  70. — Transverse  Section  through  Muscular  Fibers  of  Human  Tongue.  The 
deeply  stained  nuclei  are  situated  at  the  inside  of  the  sarcolemma.  Each  muscle  fiber 
shows  "Cohnheim's  fields,"  that  is,  the  sarcous  elements  in  transverse  section  separated 
by  clear  (apparently  linear)  interstitial  substance.  X  450.  (Klein  and  Noble  Smith.) 

mononucleated  cells,  7  to  S/j.  in  diameter  by  40  to  2Oo/z  in  length,  figures 
68  and  69.  The  protoplasm  of  each  cell,  the  contractile  substance,  is 
marked  by  longitudinal  striations  representing  fibrils  which  have  been 
described  as  contractile.  The  nucleus  is  an  oblong  mass  placed  near  the 


FIG.  71.  FIG.  72. 

FIG.  71. — Muscle  Fiber  Torn  Across;  the  sarcolemma  still  connects  the  two  parts  of  the 
fiber.  (Todd  and  Bowman.) 

FIG.  72. — Part  of  a  Striped  Muscle  Fiber  of  a  Water  Beetle  prepared  with  Absolute 
Alcohol.  At  Sarcolemma;  B,  Krause's  membrane.  The  sarcolemma  shows  regular 
bulgings.  Above  and  below  Krause's  membrane  are  seen  the  transparent  "lateral  discs." 
The  chief  mass  of  a  muscular  compartment  is  occupied  by  the  contractile  disc  composed 
of  sarcous  elements.  The  substance  of  the  individual  sarcous  elements  has  collected  more 
at  the  extremity  than  in  the  center  hence  this  latter  is  more  transparent.  The  optical 
effect  is  that  the  contractile  disc  appears  to  possess  a  "median  disc"  (Disc  of  Hensen). 
Several  nuclei,  C  and  D,  are  shown,  and  in  them  a  minute  network.  X  300.  (Klein  and 
Noble  Smith.) 

center  of  the  cell.     It  is  covered  by  a  nuclear  membrane  which  encloses  a 
network  of  anastomosing  fibrils. 

Development. — In  the  pig  the  smooth  muscle  of  the  alimentary  canal 
originates  in  the  syncytium  of  the  mesodermal  cells  which  surround  the 


60  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 

entoderm.  The  cells  soon  begin  to  grow  into  the  adult  spindle-shaped  form 
and  the  fibrils  make  their  appearance.  Even  in  the  adult  muscle  the  syn- 
cytial  connections  are  retained,  according  to  Dr.  McGill. 

STRIATED  MUSCLE. 

Striated  or  striped  muscle  constitutes  the  whole  of  the  muscular  apparatus 
of  the  skeleton,  of  the  walls  of  the  abdomen,  the  limbs,  etc. — the  whole 
of  those  muscles  which  are  under  the  control  of  the  will  and  hence  termed 
voluntary;  also  the  muscle  of  the  heart. 

For  the  sake  of  description,  striated  muscular  tissue  may  be  divided 
into  two  classes,  (a)  skeletal,  which  comprises  the  whole  of  the  striated  mus- 
cles of  the  body  except  (b)  the  heart. 


FIG.  73. — A,  Portion  of  a  Medium-sized  Human  Muscle  Fiber.  B,  Separated  bundles 
of  fibrillae  equally  magnified;  a,  a,  larger,  and  b,  b,  smaller  collections;  c,  still  smaller;  d,  d, 
the  smallest  which  could  be  detached,  possibly  representing  a  single  series  of  sarcous 
elements.  X  800.  (Sharpey.) 

Skeletal  Muscle. — The  muscle  fibers  of  the  skeletal  muscles  are  usually 
grouped  in  small  parallel  bundles,  fasciculi.  The  fasciculi  extend  through 
the  muscle,  converging  to  their  tendinous  insertions.  Connective-tissue 
sheaths,  endomysium,  surround  the  fasciculi  and  support  the  blood  vessels, 
while  a  stronger  sheath,  the  perimysium,  encases  the  entire  muscle. 

The  unit  of  muscular  structure  is  the  fiber.  Each  muscle  fiber  is  a  long 
cylinder  with  fusiform  ends.  The  fibers  vary  in  diameter  from  10  to  ioo/z, 
while  the  length  may  reach  as  much  as  40  mm.  Each  fiber  is  enclosed  in 


SKELETAL    MUSCLE 


6l 


a  distinct  sheath,  the  sarcolemma.    The  sarcolemma  is  a  transparent  structure- 
less sheath  of  great  resistance  which  surrounds  each  fiber,  figure  71. 

The  substance  of  the  fiber  enclosed 
by  the  sarcolemma,  the  contractile 
substance,  contains  a  number  of  oval 
nuclei  distributed  along  the  length  of 
the  fiber  and  lying  just  under  the 
sarcolemma  or  through  the  sarco- 
plasm.  Each  nucleus  is  accompanied 
by  a  small  mass  of  granular  proto- 
plasm at  its  poles.  The  main  mass  of 
the  fiber  is  characterized  by  transverse 
light  and  dark  bands,  figure  73,  from 
which  the  name  striated  muscle  arises. 

Longitudinal  striation  is  also  ap- 
parent under  certain  modes  of  treat- 
ment, figure  81.      The  muscle  fibers 
can  be  split  longitudinally  into  fibrils, 
called  sarcostyles,  figures  73  and  74, 
each  of  which  exhibits  the  character- 
istic   striation    of    the    whole    fiber. 
Under   certain   treatment   the   sarco- 
styles  break    transversely    into    smaller   discs   by  cleavage  at  the  line  of 
Krause's   membrane. 

The  sarcostyle  is,  therefore,  composed  of  a  number  of  smaller  elements 


8.E. 


FIG.  74. — Diagram  of  Segment  of  Muscle 
Fiber,  showing  Sarcostyle  A,  Sarcous 
element  ZJ,  Krause's  line  C,  Hensen's 
line  D. 


S.E. 


FIG.  75. 

FIG.  75. — Sarcostyles  from  the  Wing  Muscles  of  a  Wasp.  A,  A',  Sarcostyles  showing 
degrees  of  retraction;  B,  a  sarcostyle  extended  with  the  sarcous  elements  separated  into  two 
parts;  C,  sarcostyles  moderately  extended  (semidiagrammatic).  (E.  A.  Schafer.) 

FIG.  76. — Diagram  of  a  Sarcomere  in  a  Moderately  Extended  Condition,  B.  K,  K, 
Krause's  membranes;  H,  plane  of  Henson;  S,  E,  poriferous  sarcous  element.  (E.  A. 
Schafer.) 

joined  end  to  end.      These  are  the  sarcous  elements  of  Bowman.     The  sar- 
cous element  has  a  highly  refractive  denser  middle  piece  surrounded  by  a 


62 


CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 


less  refractive  more  fluid  material.  The  polarizing  microscope  reveals  the 
fact  that  the  middle  piece  which  corresponds  in  position  to  the  dark 
transverse  band  is  doubly  refractive,  anisotropic,  while  the  surrounding 
material,  the  light  band,  is  singly  refractive,  isotropic. 

In  transverse  sectioti,  figure  70,  the  area  of  the  muscle  substance  is 
mapped  out  into  small  polygonal  areas  by  a  network  of  clear  lines  called  Cohn- 
heim's  areas.  The  lines  represent  the  substance  between  the  sarcostyles. 
This  substance  probably  represents  the  less  differentiated  contractile  sub- 

stance,   called  sarcoplasm.      In  figure  81 

•  the  interfibrillar  sarcoplasm  is  indicated 

•  by  the  longitudinal  and  transverse  lines. 

Heart   Muscle. — The    muscle    sub- 
stance   of    the    heart    is    composed    of 


FIG.  77.  FIG.  78. 

FIG.  77. — A  Section  of  Cardiac  Muscle,  Diagrammatic.  (From  E.  A.  Schafer,  after 
Heidenhain.) 

FIG.  78. — Intercellular  Continuity  of  Muscle  Fibrils  in  Cardiac  Muscle.  (From  E.  A. 
Schafer  after  Przewosky.) 

mononucleated  masses  of  protoplasm,  cardiac  muscle  cells,  in  which  the 
substance  of  the  cell  presents  the  transversely  striated  appearance  char- 
acteristic of  the  voluntary  muscle  just  described.  But  the  heart  muscle  is 
physiologically  much  more  like  an  involuntary  muscle.  The  cells  are  rather 
small,  two  to  four  times  as  long  as  thick,  and  the  nucleus  is  usually  situated 
near  the  middle  of  the  cell,  figure  79.  There  is  no  sarcolemma;  on  the  other 
hand,  the  cells  present  branched  and  irregular  outlines,  but  adjacent  cells 
interlock  in  close-fitting  contact. 

Certain  observers  have  described  fibrils  as  extending  across  the  so-called 
cell  boundary  and  noted  that  not  all  such  boundaries  enclose  nuclei.  These 


BLOOD  AND  NERVE  SUPPLY  63 

observations  suggest  that  cardiac  muscle  belongs  to  the  group  of  tissues 
possessing  a  syncytium.  However,  the  section  of  cardiac  tissue  may  very 
possibly  cut  many  cells  without  enclosing  a  nucleus.  The  continuity  of 
fibrils  is  an  important  observation  from  the  physiological  point  of  view;  see 
Circulation  chapter. 

In  certain  parts  of  the  heart,  the  cardiac  tissue  is  not  completely  differ- 
entiated and  retains  in  the  adult  somewhat  embryonic  characters;  for  ex- 
ample, the  bundle  of  His  running  in  the  septum  from  the  auricles  to  the 
ventricles  and  the  cells  containing  Purkinje's  fibers  lying  immediately  under 
the  endocardium. 


FIG.  79. 


FIG.  80. 


FIG.  79.— Muscular  Fiber  Cells  from  the  Heart.     (E.  A.  Schafer.) 

FIG.  80. — From  a  Preparation  of  the  Nerve  Termination  in  the  Muscular  Fibers  of  a 

Snake,     a,  End-plate    seen    only  broad-surfaced;  6,  end-plate  seen  as  narrow  surface. 

(Lingard  and  Klein.) 


Blood  and  Nerve  Supply. — The  muscles  are  freely  supplied  with  blood 
vessels;  the  capillaries  form  a  network  with  oblong  meshes  around  the  fibers. 
Nerves  also  are  supplied  freely  to  muscles;  the  striated  voluntary  muscles 
receiving  them  from  the  cerebro- spinal  nerves,  and  the  cardiac  muscle  from 
both  the  cerebro-spinal  and  the  sympathetic  nerves. 

In  striped  muscle  the  nerves  end  in  motor  end-plates.  The  nerve  fibers 
are  medullated;  and  when  a  branch  passes  to  a  muscle  fiber,  its  primitive 
sheath  becomes  continuous  with  the  sarcolemma,  and  the  axis-cylinder 
forms  a  network  of  its  fibrils  on  the  surface  of  the  muscle  fiber.  This  net- 
work lies  embedded  in  a  flattened  granular  mass  containing  nuclei  of  several 
kinds;  this  is  the  motor  end-plate,  figures  80  and  81.  There  is  considerable 
variation  in  the  exact  form  of  the  nerve  end-plate  in  the  muscle.  In  batrachia 
the  nerve  fiber  ends  in  a  brush  of  branching  nerve  fibrils  which  are  accom- 
panied here  and  there  by  attached  oval  nuclei. 


CELL    DIFFERENTIATION   AND    THE    ELEMENTARY    TISSUES 


Development. — The  striated  muscle  of  the  voluntary  variety  is  usually 
developed  from  the  mesoderm.  The  embryonic  cells  increase  enormously 
in  size,  the  nuclei  multiply  by  fission  and  distribute  themselves  beneath  the 


FIG.  81. 


FIG.  82. 


FIG.  81. — Two  Striped  Muscle  Fibers  of  the  Hyoglossus  of  Frog,  a,  Nerve  end-plate; 
by  nerve  fibers  leaving  the  end-plate;  c,  nerve-fibers  terminating  after  dividing  into  branches; 
d,  a  nucleus  in  which  two  nerve  fibers  anastomose.  X  600.  (Arndt.) 

FIG.  82. — Developing  Striated  Muscular  Fibers,  Showing  Different  Stages  of  Develop- 
ment and  Different  Positions  of  the  Unstriated  Protoplasm.  A. — Elongated  cell  with 
two  nuclei;  the  longitudinal  striation  is  beginning  to  show  on  the  right  side.  From  a  fetal 
sheep.  (Wilson  Fox.)  B. — Developing  muscular  fiber,  showing  both  longitudinal 
and  transverse  striations  at  the  periphery,  and  a  central  unstriated  cylinder  of  protoplasm 
containing  several  nuclei.  From  a  human  fetus  near  the  third  month.  (Ranvier.)  n, 
Nucleus  (there  is  usually  a  mass  of  glycogen  near  each  nucleus);  p,  central  unstriated 
protoplasm;  s,  peripheral  striated  substance.  C. — Developing  muscular  fiber,  showing  a 
lateral  position  of  the  unstriated  protoplasm.  From  a  three  months'  human  fetus. 
(Ranvier.)  n,  Nucleus;  g,  unstriated  protoplasm  at  one  side  of  the  fiber;  s,  striated  sarcous 
substance  with  longitudinal  and  transverse  striations. 

sarcolemma.  There  is  a  differentiation  of  the  cell  protoplasm  which  takes 
place  by  the  formation  of  sarcostyles.  This  begins  nearest  the  surface  of 
the  cells  and  proceeds  toward  the  center  of  the  mass. 


NERVOUS    TISSUE  65 

The  sarcolemma  is  apparently  produced  from  embryonic  connective 
tissue. 

The  cardiac  muscle  cells  are  at  first  spindle-shaped  embryonic  cells 
which  elongate  more  and  more.  In  further  differentiation  their  protoplasm 
exhibits  faint  striations  which  pervade  the  cell  as  it  grows  in  the  great  increase 
in  size.  The  rhythmic  contractions  begin  long  before  the  striations  appear. 

IV.  NERVOUS  TISSUE. 

Nervous  tissue  has  usually  been  described  as  being  composed  of  two 
distinct  substances,  nerve  fibers  and  nerve  cells.  The  modern  view  of  the 
nature  of  nerve  tissue  is,  however,  that  the  nerve  cell  and  the  nerve  fibers 
are  to  be  considered  together  as  one  unit,  called  the  neurone.  The  neurone 


/S.N. 


FIG.  83. — Diagram  Showing  the  Arrangement  of  the  Neurones  or  Nerve  Units  in  the 
Architecture  of  the  Nervous  System.  (Raymon  y  Cajal.)  A,  Pyramidal  neurone  of 
cerebral  cortex;  B,  anterior  horn  motor  cell  of  spinal  cord;  D,  collateral  branches  of  A;  E, 
medullary  neurone  with  ascending  axone;  F,  spinal-ganglion  neurones;  G,  sensory  axones 
of  F;  7,  collaterals  of  F  in  the  cord. 

is  embedded  in,  and  supported  by,  a  substance  called  neuroglia.  This  neurone 
consists  of  a  cell  body,  a  number  of  branching  processes  termed  dendrites, 
and  a  long  process  running  out  from  it,  the  neuraxone,  or  axone,  which  be- 
comes eventually  a  nerve  fiber.  The  nerve  cell  and  the  nerve  fiber  are  parts 
of  the  same  anatomical  unit,  and  the  nervous  centers  are  made  up  of  those 
units,  arranged  in  different  ways  throughout  the  nervous  system,  figure  83. 
5 


66 


CELL   DIFFERENTIATION   AND    THE    ELEMENTARY    TISSUES 


C- 


NERVE    FIBERS. 


While  the  nerve  fiber  is  really  to  be  considered 
as  a  process  of  the  nerve  cell,  it  is  convenient  to  de- 
scribe it  separately.  Nerve  fibers  are  of  two  kinds, 
medullated  or  white  fibers,  and  non-medullated  or 
gray  fibers. 

Medullated  Fibers. — Each  medullated  nerve 
fiber  is  made  up  of  the  following  parts:  An  ex- 
ternal sheath,  called  the  primitive  sheath,  neuri- 
lemma,  or  nucleated  sheath  of  Schwann;  an  inter- 
mediate, known  as  the  medullary  or  myelin  sheath, 
or  white  substance  of  Schwann;  and  a  central 
thread,  the  axis- cylinder,  or  axial  fiber. 

The  Primitive  Sheath. — This  is  a  pellucid  mem- 
brane forming  the  outer  investment  of  the  nerve 
fiber.  The  sheath  is  constricted  at  intervals  of  a 
millimeter  or  less,  the  nodes  of  Ranvier.  Each  in- 
ternodal  segment  bears  a  single  nucleus  surrounded  by  a  variable  amount 
of  protoplasm.  This  membrane  is  described  as  having  its  origin  in  the 


FIG.  84. — Two  Nerve 
Fibers  of  the  Sciatic 
Nerve.  A,  Node  of  Ran- 
vier; B,  axis-cylinder; 
C,  sheath  of  Schwann 
with  nuclei.  X  300. 
(Klein  and  Noble 
Smith.) 


FIG.  85. — A  Node  of  Ranvier  in  a  Medullated  Nerve  Fiber,  viewed  from  above.  The 
medullary  sheath  is  interrupted  and  the  primitive  sheath  thickened.  Copied  from  Axel 
Key  and  Retzius.  X  750.  (Klein  and  Noble  Smith.) 


FIG.  86. — Gray,  Pale,  or  Gelatinous  Nerve  Fibers.  A,  From  a  branch  of  the  olfactory 
nerve  of  the  sheep;  two  dark-bordered  or  white  fibers  from  the  fifth  pair  are  associated 
with  the  pale  olfactory  fibers,  B,  from  the  sympathetic  nerve.  X  450.  (Max  Schultze.) 

mesoblastic  cells,  and  the  nuclei  are  the  indications  of  the  cellular  nature 
of  each  nodal  segment. 

The  Medullary  or  Myelin  Sheath. — This  is  the  part  to  which  the  peculiar 


NON-MEDULLATED    FIBERS  67 

opaque  white  aspect  of  medullated  nerves  is  due.  The  thickness  of  this 
layer  of  nerve  fiber  varies  considerably.  It  is  a  semifluid,  fatty  substance 
of  high  refractive  power.  It  possesses  a  fine  reticulum  (Stilling,  Klein),  in 
the  meshes  of  which  is  embedded  the  fatty  material.  It  stains  well  with 
osmic  acid. 

The  Axis-cylinder. — The  central  thread  of  a  medullated  nerve  fiber  is 
the  axis-cylinder.  It  is  the  prolongation  of  a  nerve  cell  and  extends  un- 
interrupted for  the  full  length  of  the  fiber.  It  consists  of  a  large  number  of 
primitive  fibrillcz,  as  shown  in  the  cornea,  where  the  axis-cylinders  of  nerves 
break  up  into  minute  fibrils  which  form  terminal  networks.  From  various 
considerations,  such  as  its  invariable  presence  and  unbroken  continuity  in 
all  nerves,  there  can  be  no  doubt  that  the  axis-cylinder  is  the  essential  con- 


FIG.  87. — Transverse  Section  of  a  Portion  of  the  Sciatic  Nerve  of  the  Rabbit,  Hardened 
in  Chromic  Acid  and  Stained  with  Picro-carmine,  to  show  medullated  fibers  in  end  view. 
X  275.  a,  Perifascicular  connective  tissue;  b,  lamellar  sheath;  e,  axis-cylinder. 

ducting  part  of  the  fiber,  the  other  parts  having  the  subsidiary  function  of 
support  and  possibly  of  insulation. 

The  size  of  the  nerve  fibers  varies,  figure  87.  The  largest  fibers  are 
found  within  the  trunks  and  branches  of  the  spinal  nerves,  in  which  the 
majority  measure  from  14/4  to  igp  in  diameter.  In  the  so-called  visceral 
or  autonomic  nerves  of  the  brain  and  spinal  cord  medullated  nerves  are 
found,  the  diameter  of  which  varies  from  i .  S/i  to  3 . 6/1.  In  the  hypoglossal 
nerve  they  are  intermediate  in  size,  and  generally  measure  7 .  2fj.  to  10. 8//. 

Non-medullated  Fibers. — The  fibers  of  the  second  kind,  figure  86, 
which  are  also  called  fibers  of  Remak,  constitute  the  principal  part  of  the 
trunk  and  branches  of  the  sympathetic  nerves,  the  whole  of  the  olfactory 


68 


CELL   DIFFERENTIATION  AND    THE    ELEMENTARY   TISSUES 


nerve,  and  are  mingled  in  various  proportions  in  the  cerebro-spinal  nerves. 
They  differ  from  the  preceding  chiefly  in  not  possessing  the  outer  layer  of 
medullary  substance;  their  contents  being  composed  exclusively  of  the  axis- 
cylinder. 


FIG.  88. — Transverse  Section  of  the  Sciatic  Nerve  of  a  Cat,  about  X  100.  It  consists 
of  bundles  (funiculi)  of  nerve  fibers  ensheathed  in  a  fibrous  supporting  capsule,  epineurium, 
A;  each  bundle  has  a  special  sheath  (not  sufficiently  marked  out  from  the  epineurium  in 
the  figure)  or  perineurium,  B,  the  nerve  fibers,  Nf;  L,  lymph  spaces;  Ar,  artery;  V,  vein; 
F,  fat.  Somewhat  diagrammatic.  (V.  D.  Harris.) 

The  non-medullated  nerves  are  only  about  one-third  to  one-half  as 
large  as  the  medullated  nerves,  they  do  not  exhibit  the  double  contour,  and 


FIG.  89. — Small  Branch  of  a  Motor  Nerve  of  the  Frog,  near  its  Termination,  Showing 
Divisions  of  the  Fibers:  a,  into  two;  b,  into  three.      X  350.     (Kolliker.) 


NERVE    COLLATERALS  69 

they  are  grayer  than  the  medullated  nerves.  The  non-medullated  fibers 
frequently  branch. 

It  is  worthy  of  note  that  in  the  fetus,  at  an  early  period  of  development, 
all  nerve  fibers  are  non-medullated. 

Nerve  Trunks. — Each  nerve  trunk  is  composed  of  a  variable  number 
of  different-sized  bundles,  funiculi,  of  nerve  fibers  which  have  a  special 
sheath,  perineurium.  The  funiculi  are  enclosed  in  a  firm  fibrous  sheath, 
epineurium;  this  sheath  also  sends  in  processes  of  connective  tissue  which 
connect  the  bundles  together.  In  the  funiculi  between  the  fibers  is  a  delicate 
supporting  tissue,  the  endoneurium.  There  are  numerous  lymph  spaces 
both  beneath  the  connective  tissue  investing  individual  nerve  fibers  and 
also  beneath  that  which  surrounds  the  funiculi. 


FIG.  90. — Terminal  Ramifications  of  a  Collateral  Branch  Belonging  to  a  Fiber  of  the 
Posterior  Column  in  the  Lumbar  Cord  of  an  Embryo  Calf. 

Bundles  of  fibers  run  together  in  the  nerve  trunk,  but  they  merely  lie 
in  approximation  to  each  other,  they  do  not  unite.  Even  when  nerves  anas- 
tomose, there  is  no  union  of  fibers,  but  only  an  interchange  of  fibers  between 
the  anastomosing  bundles.  Although  each  nerve  fiber  is  thus  single  through 
most  of  its  course,  yet,  as  it  approaches  the  region  in  which  it  terminates,  it 
may  break  up  into  several  subdivisions  before  its  final  ending. 

Nerve  Collaterals. — It  has  been  discovered  through  the  researches  of 
Golgi,  and  confirmed  by  the  further  studies  of  Cajal  and  other  anatomists, 


70  CELL   DIFFERENTIATION  AND    THE   ELEMENTARY   TISSUES 

that  each  individual  nerve  fiber  in  the  central  nervous  system  gives  off  in  its 
course  branches  which  pass  out  from  it  at  right  angles  for  a  short  distance, 
and  then  may  run  in  various  directions.  These  branches  are  called  collaterals. 
They  end  in  fine,  brush-like  terminations  known  as  end-brushes,  or  in  little 


FIG.  91. 


FIG.  92. 


FIG.  91. — Nerve  Cell  with  Short  Axis-cylinder  from  the  Posterior  Horn  of  the  Lumbar 
Cord  of  a  o .  55  cm.  Embryo  Calf.  (After  Van  Gehuchten.) 

FIG.  92. — Scheme  of  Lower  Motor  Neurone.  The  cell  body,  protoplasmic  processes, 
axone,  collaterals,  and  terminal  arborizations  in  muscle  are  all  seen  to  be  parts  of  a  single 
cell  and  together  constitute  the  neurone,  c,  Cytoplasm  of  cell  body  containing  chromo- 
philic  bodies,  neurofibrils,  and  perinbrillar  substance;  «',  nucleus;  n,  nucleolus;  d,  dendrites; 
ah,  axone  hill  free  from  chromophilic  bodies;  ax,  axone;  sf,  side  fibril  (collateral);  m,  medul- 
lary sheath;  nR,  node  of  Ranvier  where  side  branch  is  given  off;  si,  neurilemmaand  incisures 
of  Schmidt;  m,  striated  muscle  fiber;  tel,  motor  end-plate.  (Barker.) 


bulbous   swellings  which    come    in   close  contact  with   other  nerve  cells, 
figures  83  and  90. 

In  the  nerve  centers,  that  is,  in  the  brain  and  spinal  cord,  the  different 
nerve  fibers  end  just  as  the  collaterals  do,  by  splitting  up  into  fine  branches 
which  form  the  end-brushes.  Collaterals  of  the  nerve  fibers  and  end-brushes 


NERVE    COLLATERALS 


are  chiefly  found  in  the  nervous  centers.  The  nerve  fibers  of  the  peripheral 
nerves  end  in  the  muscles,  glands,  or  special  sensory  organs,  such  as  the 
eye  and  ear,  each  by  its  own  special  type  of  ending.  Here,  however,  some 


FIG.  93. — Large  Nerve  Cells  with  Processes,  from  the  Ventral  Cornua  of  the  Cord  of 
Man.  X  350.  On  the  cell  at  the  right  two  short  processes  of  the  cell  body  are  present, 
one  or  the  other  of  which  may  have  been  an  axis-cylinder  process  (Deiters).  A  similar 
process  appears  also  on  the  cell  at  the  left. 


FIG.  94. — Multipolar  Nerve  Cell  of  the  Cord  of  an  Embryo  Calf. 

analogy  to  the  end-brush  can  also  be  discovered.  As  the  peripheral  nerve 
fibers  approach  their  terminations,  they  lose  their  medullary  sheath,  and 
consist  then  merely  of  an  axis-cylinder  and  primitive  sheath.  They  may 
even  lose  the  latter,  and  only  the  axis-cylinder  be  left.  Finally,  the  axis- 


72  CELL   DIFFERENTIATION  AND   THE   ELEMENTARY   TISSUES 

cylinder  breaks  up  into  its  elementary  fibrillae,  to  end  in  various  ways  to 
be  described  later. 


THE  NERVE -CELL  BODY. 

The  nerve-cell  body  is  the  nodal  and  important  part  of  the  neurone,  and 
from  it  are  given  off  the  dendrites  and  axis-cylinder  process  or  axone.  It 
consists  of  a  mass  of  protoplasm,  of  varying  shape  and  size,  containing  within 
it  a  nucleus  and  nucleolus.  All  nerve  cells  give  off  one  or  more  processes 
which  branch  out  in  various  directions,  dividing  and  subdividing  like  the 
branches  of  a  tree,  but  never  anastomosing  with  each  other  or  with  other  cells. 


FIG.  95. — Ganglion  Cells,  Showing  Neurofibrils.     A,  Anterior-horn  cells  of  human;  B,  cell 
from  the  facial  nucleus  of  rabbit;  C,  dendrite  of  anterior  horn  cell  of  human.     (Bethe.) 

These  branches  are  what  have  already  been  referred  to  as  the  dendrites  of 
the  cell.  They  were  formerly  called  the  protoplasmic  processes,  figures  91, 
93.  It  is  thus  seen  that  the  neurone  or  nerve  unit  consists  of  a  number  of 
subdivisions,  namely,  the  cell  body,  with  its  nucleus  and  nucleolus,  the 
dendrites  or  protoplasm  processes,  and  the  axone  or  axis-cylinder  process. 

The  protoplasm  of  the  cells  is  shown  by  various  dyes  to  consist  of  neuro- 
fibrils,  periftbrillar  substance,  and  in  most  cells  chromophilic  bodies.  Apathy 
and  others  have  demonstrated  that  a  network  of  interlacing  and  anasto- 
mosing fibrils  traverses  both  the  cell  body  and  its  branches,  figure  95. 

The  perifibrillar  substance  is  a  fluid  or  semifluid  substance  in  which  the 
fibrils  are  embedded.  By  treating  nerve  cells  with  special  stains  granular 
bodies  varying  in  size  are  found  embedded  in  the  cytoplasm.  These  bodies 
are  the  chromophilic  bodies,  figure  96. 


THE    NERVE-CELL    BODY 


73 


Ganglion  cells  are  generally  enclosed  in  a  transparent  membranous 
capsule  similar  in  appearance  to  the  external  nucleated  sheath  of  nerve 
fibers;  within  this  capsule  is  a  layer  of  small  flattened  cells. 


FIG.  96. — Cell  of  the  Anterior  Horn  of  the  Human  Spinal  Cord,  Stained  by  NissPs  Method, 
showing  chromophiles  in  blue,  and  pigment  in  black.     (After  Edinger.) 


c- 


FIG.  97. — An  Isolated  Sympathetic  Ganglion  Cell  of  Man,  Showing  Sheath  with 
Nucleated  Cell  Lining,  B.  A,  Ganglion  cell,  with  nucleus  and  nucleolus;  C,  branched 
process  or  dendrite;  D,  unbranched  process  or  axone.  (Key  and  Retzius.)  X  750. 


74  CELL   DIFFERENTIATION  AND   THE   ELEMENTARY   TISSUES 

Nerve  Terminations. 

Nerve  fibers  terminate  peripherally  in  four  different  ways:  i,  by  the  ter- 
minal subdivisions  which  pass  in  between  epithelial  cells,  and  are  known  as 
interepithelial  arborizations;  2,  by  motor-plates  which  lie  in  the  muscles; 
3,  by  special  end-organs,  connected  with  the  sense  of  sight,  hearing,  smell, 
and  taste;  and  4,  by  various  forms  of  tactile  corpuscles. 


P'iG.  98. — Sensory  Nerve  Terminations  in  Stratified  Pavement  Epithelium.     Golgi's  rapid 
method.     (After  G.  Retzius.) 

The  Interepithelial  Arborizations. — This  forms  a  most  common 
mode  of  termination  of  the  sensory  nerves  of  the  body.  The  nerve  fibers 
to  the  surface  of  the  skin  or  mucous  membrane  lose  their  neurilemmse  and 
myelin  sheaths,  the  bare  axis-cylinder  divides  and  subdivides  into  minute 
ramifications  among  the  epithelial  cells  of  the  skin  and  mucous  membrane. 
In  the  various  glands  of  the  body  this  form  of  termination  also  prevails. 
The  hair  bulbs,  the  teeth,  and  the  tendons  of  the  body  are  supplied  by  this 
same  process  of  terminal  arborization,  figures  98,  99. 


FIG.  99. — Sensory  Nerve  Termination  in  the  Epithelium  of  the  Mucosa  of  the  Inferior 
Vocal  Cord  and  in  the  Ciliated  Epithelium  of  the  Subglottic  Region  of  the  Larynx  of  a 
Cat  Four  Weeks  Old.  Golgi's  rapid  method,  n,  Nerve  fibers  rising  from  the  connective- 
tissue  layer  into  the  epithelial  layer,  where  they  terminate  in  ramified  and  free  arborizations. 
(After  G.  Retzius.) 

The  motor  nerves  to  the  muscles  end  in  what  are  known  as  muscle  plates, 
the  details  of  whose  structure  have  been  already  described. 

The  special  sensory  end-organs  will  be  described  later  in  the  chapter 
on  the  Special  Senses. 

A  fourth  form  of  termination  consists  of  corpuscles  that  are  more  or  less 
encapsulated,  and  these  are  known  as  the  corpuscles  of  Pacini,  the  tactile 


THE    TACTILE    CORPUSCLES    OF    MEISSNER 


75 


corpuscles  cf  Meissner,  the  tactile  corpuscles  of  Krause,  the  tactile  menisques, 
and  the  corpuscles  of  Golgi. 

The  Pacinian  Corpuscles. — These  nerve  endings,  named  after  their 
discoverer  Pacini,  are  elongated  oval  bodies  situated  on  some  of  the  cerebro- 
spinal  and  sympathetic  nerves.  They  occur  on  the  cutaneous  nerves 
of  the  hands  and  feet,  the  branches  of  the  large  sympathetic  plexus  about 
the  abdominal  aorta,  the  nerves  of  the  mesentery,  and  have  been  observed 
also  in  the  pancreas,  lymphatic  glands,  and  thyroid  glands,  figure  100. 
Each  corpuscle  is  attached  by  a  narrow  pedicle  to  the  nerve  on  which  it  is 


FIG. 


FIG.  ioi. 


FIG.  100. — Pacinian  Corpuscle  of  the  Cat's  Mesentery.  The  stalk  consists  of  a  nerve 
fiber,  n,  with  its  thick  outer  sheath.  The  peripheral  capsules  of  the  Pacinian  corpuscle  are 
continuous  with  the  outer  sheath  of  the  stalk.  The  intermediary  part  becomes  much 
narrower  near  the  entrance  of  the  axis-cylinder  into  the  clear  central  mass.  A  hook- 
shaped  termination  with  the  end-bulb,  a,  is  seen  in  the  upper  part.  (Ranvier.) 

FIG.  ioi. — Summit  of  a  Pacinian  Corpuscle  of  the  Human  Finger,  showing  the 
Endothelial  Membranes  Lining  the  Capsules.  X  220.  (Klein  and  Noble  Smith.) 


situated,  and  is  formed  of  several  concentric  layers  of  fine  membrane,  each 
layer  being  lined  by  endothelium,  figure  ioi.  A  single  nerve  fiber  passes 
through  its  pedicle,  traverses  the  several  concentric  layers,  enters  a  central 
cavity,  and  terminates  in  a  knob-like  enlargement  or  in  a  bifurcation. 

The  physiological  import  of  these  bodies  is  still  obscure. 

The  Tactile  Corpuscles  of  Meissner. — They  are  found  in  the  papillae 
of  the  skin  of  the  fingers  and  toes  or  among  its  epithelium.  When  simple 
they  are  small,  slightly  flattened  transparent  bodies  composed  of  nucleated 
cells  enclosed  in  a  capsule.  When  compound,  the  capsule  contains  several 


76 


CELL   DIFFERENTIATION  AND    THE   ELEMENTARY   TISSUES 


small  cells.  The  nerve  fiber  penetrates  the  corpuscles,  loses  its  myelin 
sheath,  and  divides  and  subdivides  to  form  a  series  of  arborizations.  The 
terminal  arborizations  occupy  the  central  part  of  the  corpuscle,  and  are 


FIG.  102. — Tactile  Corpuscle  of  Meissner,  Tactile  Cell,  and  Free  Nerve  Ending, 
a,  Corpuscle  proper,  outside  of  which  is  seen  the  connective-tissue  capsule;  b,  fiber 
ending  on  tactile  cell;  c,  fiber  ending  freely  among  the  epithelial  cells.  (Merkel-Henle.) 

surrounded  by  a  great  number  of  marginal  cells.     The  tactile  corpuscles 
of  Meissner  serve  for  the  special  purpose  of  touch. 

The  Corpuscles   of    Krause   or  End-bulbs. — These  exist  in  great 


FIG.  103.  FIG.  104. 

FIG.  103. — End-bulb  of  Krause.     a,  Medullated  nerve  fiber;  b,  capsule  of  corpuscle. 
FIG.  104. — A  Termination  of  a  Medullated  Nerve  Fiber  in  Tendon,  lower  half  with 
Convoluted  Medullated  Nerve  Fiber.     (Golgi.) 

numbers  in  the  conjunctiva,  the  glans  penis,  clitoris,  lips,  skin,  and  in  tendon 
of  man.  They  resemble  the  corpuscles  of  Pacini,  but  have  much  fewer 
concentric  layers  to  the  corpuscle,  and  contain  a  relatively  voluminous  central 


THE    CORPUSCLES    OF    GOLGI. 


77 


mass  composed  of  polyhedral  cells.  In  man  these  corpuscles  are  spherical 
in  shape,  and  receive  many  nerve  fibers  which  wind  through  the  corpuscles 
and  end  in  the  free  extremities,  figure  103. 

Tactile  Menisques. — In  different  regions  of  the  skin  of  man,  one  meets, 
in  the  superficial  layers  and  in  the  Malpighian  layers,  nerves  which,  after 
having  lost  their  myelin  sheath,  divide  and  subdivide  to  form  extremely 
beautiful  arborizations.  The  branches  of  these  arborizations  are  the  tactile 
menisques.  These  menisques,  which  simulate  the  form  of  a  leaf,  represent  a 
mode  of  terminal  nervous  arborization  (Ranvier). 

The  Corpuscles  of  Golgi. — These  are  small  terminal  plaques  placed 
at  the  union  of  tendons  and  muscles,  but  belonging  more  properly  to  the 


FIG.  105. — Neuroglia  Cells  in  the  Cord  of  an  Adult  Frog.  A,  Ependyma  cells 
with  their  peripheral  extremities  atrophied  and  ramified;  B,  C,  D,  neuroglia  cells  in  different 
degrees  of  emigration  and  separation  from  the  ependymal  canal;  their  central  extremity 
is  atrophied  and  much  contracted;  their  peripheral  extremity,  on  the  other  hand,  is 
greatly  extended;  the  ramifications  of  the  latter,  terminating  in  conical  buttons,  7,  end 
under  the  pia  mater.  (After  Cl.  Sala.) 


tendon.  They  are  fusiform  in  shape  and  are  flattened  upon  the  surface  of 
the  tendon  close  to  its  insertion  into  the  muscular  fibers.  They  are  composed 
of  a  granular  substance,  enveloped  in  several  concentric  hyaline  membranes 
which  contain  some  nuclei.  The  nerve  fiber  passes  into  this  little  corpuscle, 
splitting  itself  up  into  fine  terminals.  The  corpuscles  of  Golgi  are  believed 
to  be  related  to  the  muscular  sense,  figure  104. 


78  CELL   DIFFERENTIATION  AND    THE    ELEMENTARY    TISSUES 

The  Muscle  Spindles. — Voluntary  muscles  are  supplied  with  nerve 
terminations  of  a  sensory  nature  ending  in  Pacinian  corpuscles,  in  end  bulbs, 
and  in  special  structures  known  as  neuromuscular  bundles  or  muscle  spindles. 
A  muscle  spindle  consists  of  one  or  more  muscle  fibers  somewhat  smaller  than 
the  typical  fibers  of  that  particular  muscle,  and  containing  a  relatively  great 
amount  of  sarcoplasm,  and  many  nuclei.  These  fibers  are  intimately  bound 
with  nerve  terminations  as  shown  in  the  figure  106.  Certain  of  the  vol- 
untary muscles,  particularly  those  of  the  arms  and  legs,  contain  large  num- 

S 


FIG.  106. — A  section  of  a  muscle  spindle  from  a.  voluntary  muscle  of  the  cat. 
A,  Annular  terminations;  S,  spiral  terminations;  F,  arborescent  terminations.  (From 
Barker,  after  Ruffini.) 

bers  of  muscle  spindles.  In  the  tendons  of  these  muscles  also  are  numerous 
muscle-tendon  organs  of  the  Golgi  type.  Sherrington  conclusively  demon- 
strated the  sensory  nature  of  these  fibers  and  terminations  by  showing  that 
they  did  not  undergo  Wallerian  degeneration  when  the  corresponding  anterior 
spinal  nerve  roots  were  cut  and  allowed  to  degenerate. 

THE  NEUROGLIA. 

The  neuroglia,  while  not  a  nervous  tissue,  is  closely  mingled  with  it  and 
forms  an  important  constituent  of  the  nervous  system.  It  consists  of  cells 
giving  off  a  fine  network  of  richly  branching  fibers.  Neuroglia  is  a  form 
of  connective  tissue,  and  it  is  in  its  functions  strictly  comparable  to  the  con- 
nective tissue  which  supports  the  special  structures  of  other  organs,  like  the 
lungs  and  kidneys.  In  the  adult  animal  the  neuroglia  tissue  is  composed  of 
cells  from  which  are  given  off  immense  numbers  of  fine  processes.  These 
extend  out  in  every  direction,  and  intertwine  among  the  nerve  fibers  and  nerve 
cells,  figure  105.  The  neuroglia  cell  differs  in  size  and  shape  very  much  in 
different  parts  of  the  nervous  system  in  accordance  with  the  arrangement  of 
the  nervous  structures  about  it.  The  cell  is  composed  of  granular  proto- 
plasm, and  lying  in  it  is  a  large  nucleus,  within  which  is  a  nucleolus.  The 
body  of  the  cell  is  small  in  proportion  to  the  nucleus. 


CHAPTER  III. 
THE  CHEMICAL  COMPOSITION  OF  THE  BODY. 

OF  the  eighty  chemical  elements  which  have  been  isolated,  no  less  than 
seventeen  combine  in  varying  quantities  to  form  the  chemical  basis  of  the 
animal  body.  The  substances  which  contribute  the  largest  share  are  the 
non- metallic  elements,  Nitrogen,  Oxygen,  Carbon,  and  Hydrogen — oxygen 
and  carbon  making  up  altogether  about  85  per  cent,  of  the  whole.  The 
most  abundant  of  the  metallic  elements  are  Calcium,  Sodium,  and  Potassium.* 

These  elements  do  not  exist  in  the  animal  body  in  the  free  state,  but  are 
combined  into  complex  chemical  compounds. 

The  first  step  in  the  act  of  separating  the  composition  products  of  proto- 
plasm produces  changes  which  destroy  the  chemical  and  physical  relations 
of  these  products  which  maintain  the  state  of  life.  Dead  protoplasm,  how- 
ever, yields  a  number  of  substances  which  must  be  very  directly  derived 
from  the  living  protoplasm.  On  the  other  hand,  certain  products  can  be 
isolated  from  the  animal  body  wrhich  are  evidently  not  a  part  of  the  proto- 
plasm itself,  but  products  of  protoplasmic  activity.  Some  of  these,  like  fat, 
glycogen,  etc.,  are  constructive  products,  others  are  disintegration  products 
of  protoplasmic  activity. 

THE  NITROGENOUS  SUBSTANCES. 

The  nitrogenous  substances  in  the  body  consist  chiefly  of  the  proteins 
or  of  substances  which  are  derived  from  the  proteins.  Nitrogenous  sub- 
stances occur  in  the  solid  tissues  of  the  body  and  are  found  also  to  a  con- 
siderable extent  in  the  circulating  fluids  (the  blood  and  lymph)  and  in  the 
secretions  and  excretions. 

*  The   following   table  represents  the  relative  proportion  of  the  various  elements  in 
the  body.     (Marshall.) 


Oxygen 72.0 

Carbon 13 . 5 

Hydrogen 9.1 

Nitrogen    2.5 

Calcium 1.3 

Phosphorus *  •  J5 

Sulphur o.  1476 

Sodium o.i 

Chlorine 0.085 

79 


Fluorine o. 08 

Potassium o .  026 

Iron o.oi 

Magnesium 0.012 

Silicon   o. 0002 

(Traces  of  copper,  lead,  and  alu- 
minum)   


100 


8o  THE   CHEMICAL   COMPOSITION    OF   THE    BODY 

THE  PROTEINS. 

These  nitrogenous  substances  constitute  the  most  important  and  com- 
plex compounds  in  the  body.  They  are  essentially  the  organic  basis  of  all 
living  substance.  At  the  same  time  they  are  the  most  important  of  our 
organic  food  stuffs.  The  proteins  are  necessary  as  food  material  for  the 
continuance  of  life  and  cannot  be  replaced  in  the  diet  by  any  other  organic 
or  inorganic  substances.  Without  them  all  life,  whether  animal  or  vegetable, 
is  impossible. 

The  proteins  are  substances  containing  carbon,  hydrogen,  and  oxygen 
(which  are  present  in  fats  and  carbohydrates).  The  proteins  also  contain 
nitrogen  and  sulphur.  Phosphorus  and  certain  metallic  elements  are  present 
as  constituents  of  some  proteins.  The  elementary  composition  of  most 
protein  substances  falls  within  the  following  percentages: 

Carbon from  50      to  55.0  per  cent. 

Hydrogen from    6      to  7 .3  per  cent. 

Oxygen    from  19      to  24.0  per  cent. 

Nitrogen    from  15      to  19.0  per  cent. 

Sulphur    from     0.3  to  2.5  per  cent. 

Phosphorus,  when  present from     0.4  to  0.8  per  cent. 

The  individual  protein  substances  are  chemical  entities.  As  individuals 
of  a  group  they  differ  in  elementary  composition  and  in  the  derivatives 
which  they  yield  on  cleavage  of  the  protein  molecule. 

Chemical  Structure  of  Proteins. — Proteins  are  combinations  of 
a-amino  acids,  the  simplest  example  of  which  is  glycocoll  or  a-amino  acetic 
acid.  Acetic  acid  has  the  formula  CH3COOH;  if  the  NH2  group  is  sub- 
stituted for  one  of  the  H's  in  the  CH3  radical  it  is  an  amino  acid.  The 
introduction  of  the  amino  group  in  this  way  yields  bodies  which  combine 
both  with  acids  and  with  bases.  It  is  also  possible  for  the  amino  acids  to 
combine  with  one  another,  with  the  elimination  of  water.  The  reaction, 
however,  can  only  be  brought  about  under  certain  conditions.  For  instance, 
glycocoll  can  be  combined  with  itself  as  a  dipeptid  or  combined  with  any 
other  amino  acid.  The  combination  may  be  indicated  by  the  following: 


CH2  -  NH2  -  CO  I  OH  +H  !  HN  -  CH2  -  COOH- 

glycocoll  plus  glycocoll 

CH2  -  NH2  -  CO  -  NH  -  CH2  -  COOH  +  H2O 

glycyl-glycine  plus  water 

A  glance  at  the  chemical  formulae  of  the  resulting  dipeptid  indicates 
that  in  this  new  substance  there  is  still  an  amino  group  (NH2)  and  a  carboxyl 
group  (COOH)  which  are  not  combined.  Another  amino  acid  may  be 
joined  on  to  the  carboxyl  root,  and  yet  a  fourth  on  to  the  remaining  amino 


THE    PROTEINS 


8l 


group,  so  that  the  still  more  complex  peptids  can  be  formed.  When  this  is 
done  there  is  yet  a  carboxyl  and  an  amino  group  to  which  other  amino  acids 
can  similarly  be  joined.  The  structure  of  the  protein  molecule  accordingly 
may  be  represented  as  follows: 


H 


H 


H 


-  OC  -  C  -  NH  -  OC  -  C  -  NH  -  OC  -  C  -  NH  - 


R 


R 


R 


In  which  R  indicates  here  the  rest  of  the  formula  for  any  of  the  a-amino 
acids  entering  into  the  constitution  of  protein. 

At  least  eighteen  amino  acids  have  been  found  to  enter  into  the  composi- 
tion of  the  proteins.  The  list  includes  glycocoll,  alanine,  serine,  phenyl- 
alanine,  tyrosine,  tryptophane,  cystine,  leucine,  isoleucine,  amino-butyric 
acid,  aspartic  acid,  glutaminic  acid,  proline,  oxyproline,  histidine,  arge- 
nine,  lysine,  and  diaminotrihydroxydodecanoic  acid.  Associated  with 
the  amino  acids  there  is  usually  a  detectable  amount  of  amino-carbohydrate. 
The  chemical  formulae  for  the  more  important  of  these  are  given  below: 


NH2 

H-C-COOH 
H 

Glycocoll. 


H     NH2 

I       I 
H -C-C-COOH 

H     H 

Alanine. 


H  NH2 

I  I 
C    -    C-C-COOH 

/S     I  i 

HC      CH    H  H 

I        I 
HC      CH 


C 
H 


Phenylalanine. 


(amino  acetic  acid.)        (a-amino  proprionic  acid.)      (Phenyl  a-amino  proprionic  acid.) 

H    NH, 

H  H 


OHNH2 

H-C-C-COOH 
H    H 


Serine. 

(a-amino  /3-hydroxy 
proprionic  acid.) 
6 


I       I 
C    -    C-C-COOH 

HC      CH   H    H  H-C-S-S-C-H 


HC      CH 
C 


Tyrosine. 

(p-oxyphenyl  a-amino 
proprionic  acid.) 


|  I 

H-C-NH2H-C-NH2 

I  I 

COOH          COOH 

Cystine. 


82 


THE   CHEMICAL   COMPOSITION   OF   THE   BODY 


H 

c 


HC 


H     NH2 

I       I 
-C-C-C-COOH 

II       I       I 
CH  H     H 


C      NH 
H 

Tryptophane. 
(Indol  amino  proprionic  acid.) 


CH3H     NH2 

I        I       I 
H- C-C-C-COOH 

I        I       I 
CH3H     H 

Leucine. 
(a-amino  isobutylacetic  acid.) 

H     NH2 

I       I 
HC=C-C-C-COOH 

I        I       I       I 
HN     N     H     H 


CH3NH2 

H-C-C-COOH 

I       I 
CH3H 


Valine. 
(a-amino  isovalerianic  acid.) 

NH, 


NH2 

H-C-COOH 

H-C-COOH                     | 

H-C-H 

H-C-COOH 

I 

H-C-COOH 

H                                       | 

H 

Asi 

tartic  acid,. 

Glutaminic  acid. 

C 
H 


(amino  succinic  acid.)     (a-amino  normal  glutaric 

acid). 


H    H    H     NH2 

I       I  I 

H  -N  -C  -C  -C  -C  -  COOH 

III  I 

NH=C     H    H     H    H 

NH, 


Histidine. 
(a-amino  /?-imidazol  proprionic  acid.) 

NH2H     H     H     NH2 


Arginine. 
(guanidine  a-amino  valerianic  acid.) 


H-C-C-C-C-C-  COOH 


H     H     H     H     H 

Lysine. 
(a-e-diamino  caproic  acid.) 

The  amino  acids  belong  either  to  what  is  known  in  organic  chemistry  as 
the  aliphatic,  carbocyclic,  or  heterocyclic  series;  that  is,  they  are  derivations 
either  of  the  hydrocarbons,  of  benzene  or  of  closed-ring  compounds  not 
composed  wholly  of  carbon  atoms,  but  in  which  one  or  more  of  the  links 
in  the  closed  chain  are  supplied  by  other  polyvalent  elements  (in  the  proteins 
by  nitrogen).  Thus  tyrosine  and  phenylalanine  are  carbocyclic  compounds; 
histidine,  proline,  and  tryptophane  are  heterocyclic  compounds,  and  the 
remaining  members  of  the  list  are  aliphatic  derivatives. 

Of  the  elements  of  the  protein  molecule,  nitrogen  is  by  far  the  most 


THE    PROTEINS 


characteristic  and  important.  The  animal  organism  is  unable  to  construct 
the  amino  acid  molecules  and  hence  cannot  build  up  protein  material  from 
nitrogen  of  the  atmosphere  or  from  combinations,  such  as  ammonia,  nitrates, 
and  nitrites.  Plants,  however,  have  the  property  of  synthesizing  proteins 
from  inorganic  nitrogen.  The  nitrogen  of  amino  acids  and  protein  is  directly 
utilizable  by  the  body,  so  that  the  animals  are  ultimately  dependent  upon 
plants  for  their  protein-supply. 

Nitrogen  in  the  protein  molecule  occurs  in  four  different  forms: 

1.  The  monamino  acid  nitrogen,  or  the  nitrogen  that  is  in  the  NH2  (amino) 
group  of  the  a  position. 

2.  The  diamino  acid  nitrogen,  or  the  basic  nitrogen,  as  shown  in  the 
amino  group  in  lysine. 

3.  Amide  nitrogen;  the  OH  of  the  second  COOH  group  in  the  dibasic 
glutaminic  and  aspartic  acids  in  protein  may  be  replaced  by  the  amino  group. 
On  cleavage,  the  NH2  is  split  off  from  the  acid  amide  as  ammonia. 

4.  The  guanidine  residue  as  in  arginine. 

The  distribution  of  nitrogen  in  the  protein  accordingly  depends  on  the 
amino  acids  entering  into  its  composition. 

Sulphur  of  the  protein  is  present  in  the  amino  acids,  cystine  and  cystein. 

It  has  been  stated  that  the  amino  acids  are  combined  together  in  a  pro- 
tein molecule,  carboxyl  with  amino  radical.  On  boiling  the  proteins  with 
mineral  acids,  the  reaction  is  reversed  and  the  protein  substances  are  split, 
with  the  combining  of  water,  into  the  individual  amino  acid  components. 
This  change  is  termed  a  "hydroly tic  cleavage."  The  qualitative  and  quan- 
titative determination  of  the  products  thus  obtained  have  shown  us  that  the 
proteins  differ  chemically  both  as  to  the  individual  amino  acids  which  enter 
into  the  complex  protein  molecule  and  the  amount  of  each  acid  present. 
Proteins,  then,  which  may  give  exactly  the  same  percentage  composition  and 
elementary  analysis  and  which  show  practically  the  same  physical  prop- 
erties are  found  to  be  actually  different  individuals  of  the  protein  group 
when  the  products  of  their  hydrolytic  cleavage  are  investigated.  For  exam- 
ple, the  following  tables  give  the  elementary  composition  and  the  amino 
acids  obtained  from  three  proteins  which  are  present  in  wheat  flour. 

ELEMENTARY  COMPOSITION  OF  WHEAT  PROTEINS. 


Gliadin 

Glutenin 

Leucosin 

Carbon  

S2.72 

C2.34 

•>3«O2 

Hydrogen  .  .  . 

6.86 

6.8* 

6.84 

Nitrogen  .  . 

17  66 

I  7.4.0 

16.80 

Sulphur 

i  03 

I  08 

1.28 

Oxygen  

2  I.  7"? 

22.26 

22.06 

100.  OO 

100.00 

IOO.OO 

84  THE   CHEMICAL  COMPOSITION   OF   THE   BODY 

AMINO  ACIDS  OBTAINED  ON  HYDROLYTIC  CLEAVAGE  OF  WHEAT  PROTEIN. 


Percent  of 

Gliadin 

Glutenin 

Leucosin 

Glycocoll 

o  oo 

o  89 

O   Q4 

Alanin  

2   OO 

4  6<C 

44  ^ 

Amino  valerianic  acid 

O   2  I 

O   24. 

o  18 

Leucine 

s  61 

5O  ^ 

I  I    3.4 

Proline  

7.06 

4.27, 

7,     l8 

Phenylalanine 

2    ~l  ? 

I  Q7 

7  ST. 

Aspartic  acid  

0.^8 

O.OI 

7.7,  C 

Glutaminic  acid  

•2  7.7  7 

27.42 

6  77. 

Serine  . 

O    I  7, 

O  74 

O     OO 

Tyrosine  

I.  2O 

4.2  < 

7,.  74 

Cystine  

O4^ 

O  O2 

O    OO 

Lysine 

O  OO 

I   O2 

2  7  e 

Histidine  

0.61 

1.76 

2.87 

Arginine  

7.  16 

4  72 

5O4 

Ammonia 

51  j 

4OI 

141 

Tryptophane  

present 

present 

present 

It  has  been  indicated  that  the  synthesis  in  very  simple  proteins  can  be 
attained  by  combining  amino  acids.  A  synthesis,  however,  can  be  brought 
about  by  the  reversible  action  of  the  digestive  enzymes.  Recently  Taylor 
has  been  able  to  synthesize  a  simple  protein,  a  protamine,  by  the  reversible 
action  of  a  trypsin  on  amino  acids  which  were  previously  obtained  by  the 
digestion  of  the  protamine.  Cleavage  by  enzymes  is  a  hydrolysis  of  the 
same  type  as  that  mentioned  above  by  the  use  of  the  mineral  acids.  In 
the  concentration  of  the  products  of  a  digestion  the  enzymes  act  in  the 
reverse  manner  and  resynthesize  these  substances  on  which  they  have  pre- 
viously acted.  Robertson  has  demonstrated  a  similar  reversible  action  of 
pepsin  on  paranuclein  derived  from  a  digestion  of  casein.  The  reversible 
action  may  be  indicated  by  the  equation: 


PROTEIN  +  WATER  +±  AMINO  ACIDS. 

These  experiments  lend  a  new  stimulus  to  the  efforts  to  build  up  proteins 
in  the  chemical  laboratory  along  the  lines  of  catalytic  action  of  enzymes. 
The  proteins  of  the  various  tissues  of  the  body  are  quite  different  in  character 
and  chemical  constitution  and  are  different  from  the  protein  of  the  diet. 
In  digestion  there  is  a  breaking  down  of  the  protein  in  the  food  into  simple 
combinations  of  amino  acids  or  the  acids  themselves,  and  subsequently 
there  is  a  selection  of  certain  amino  acids  and  resynthesis  of  characteristic 
proteins  by  the  cells  of  the  tissues. 


THE    PROTEINS  85 

It  is  customary  to  assign  to  a  compound  having  an  unknown  molecular 
mass,  i.e.,  relative  weight  in  units  of  the  weight  of  an  atom  of  hydrogen,  a 
formula  representing  the  least  mass  which  the  substance  could  have  and 
preserve  its  characteristic  properties.  The  simplest  formula  for  oxyhemoglo- 
bin,  the  compound  protein  of  the  red  blood  cells,  is  C658H1181N207S2FeO210. 
This  formula  is  based  on  the  relative  proportion  that  the  C,  H,  N,  S, 
and  O  bear  to  the  Fe  as  determined  by  analysis.  This  protein  contains 
iron,  and  the  least  Fe  that  one  molecule  can  contain  is  one  atom.  By  addi- 
tion of  the  atomic  masses  of  the  total  number  of  atoms  of  each  element,  the 
least  possible  molecular  mass  for  oxyhemoglobin  is  about  15,000.  It  might 
just  as  well  be  30,000  with  two  atoms  of  iron  in  the  compound.  The  follow- 
ing formulae  have  been  proposed  for  ovalbumin  and  seralbumin: 

Ovalbumin,  C239H386N58S2O78 
Seralbumin,  C450H720N116S6O140 

the  molecular  masses  being  in  the  neighborhood  of  5000  and  6000,  re- 
spectively. 

Besides  the  amino  acids  other  radicals  are  present  in  some  proteins.  A 
carbohydrate  moiety  is  evidently  present  in  certain  proteins  and  phosphoric 
acid  in  others  (as  in  the  milk  protein  casein).  Such  proteins  are  to  be  dis- 
tinguished from  those  which  exist  as  combinations  with  definite  chemical 
entities,  as  hematin,  nucleic  acid,  amino  sugars,  lecithins,  etc. 

Properties  of  Protein. — Many  proteins  have  been  prepared  in  crys- 
talline form,  especially  the  reserve  proteins  from  various  seeds.  Very  few 
animal  proteins  have  ever  as  yet  been  crystallized — seralbumin,  lactal- 
bumin,  ovalbumin  and  hemoglobin  are  examples. 

The  majority  of  the  proteins  are  soluble  in  water  or  in  dilute  solutions 
of  neutral  salts  of  strong  bases  with  alkalies.  The  proteins  do  not  form 
solutions  as  do,  for  instance,  the  inorganic  salts,  but  are  to  be  regarded  as  a 
suspension  of  the  molecules  or  molecular  aggregates.  Such  a  solution  is 
known  as  a  colloidal  solution,  and  the  proteins  are  frequently  spoken  of  as 
colloids.  Colloidal  solutions  of  the  heavy  metals  can  be  formed  by  the 
interrupted  contact  of  metal  electrodes  under  water.  The  metallic  colloidal 
solutions  and  the  protein  solutions  have  many  properties  in  common. 

When  a  true  solution  of  a  chemical  substance  of  relatively  small  molecular 
weight  is  placed  within  a  parchment  or  animal  membrane,  and  the  whole 
immersed  in  water,  the  substance  in  solution  will  diffuse  through  the  pores 
of  the  membrane  into  the  water  external  to  it;  similarly,  water  will  pass 
through  the  membrane  to  the  interior.  After  some  time  the  system  will 
come  into  equilibrium.  The  force  which  drives  the  dissolved  substance  from 
the  more  concentrated  to  the  less  concentrated  solution  is  known  as  osmotic 
pressure.  The  large  protein  molecules  and  molecular  aggregates  cannot 
pass  through  the  pores  of  the  membrane,  or,  in  other  words,  they  are  not 


86  THE    CHEMICAL   COMPOSITION    OF    THE    BODY 

diffusible.  Their  osmotic  pressure  is  very  slight.  Some  of  the  simpler 
derived  proteins,  however,  are  diffusible. 

Proteins  in  aqueous  solution  rotate  polarized  light  to  the  left.  The 
specific  rotation  of  the  individual  proteins  varies.  The  compound  proteins, 
hemoglobin  and  nucleoproteins  are  dextrorotatory. 

Proteins  chemically  are  rather  unstable  bodies  and  are  easily  hydrolyzed, 
through  heating  and  standing  in  alcohol,  into  modifications  which  differ  only 
slightly  from  the  original  substances.  This  change  is  ordinarily  termed 
coagulation. 

Proteins  may  be  precipitated  from  their  solutions  by  the  addition  of  the 
neutral  inorganic  salts  in  high  concentration  or  to  saturation.  This  pre- 
cipitation is  essentially  a  physical  one,  the  protein  remaining  unchanged. 
Salting  out  is,  then,  a  most  valuable  method  for  the  separation  and  purifica- 
tion of  the  protein  substances.  The  proteins  form  insoluble  combinations 
with  the  so-called  alkaloidal  reagents;  phosphotungstic  acid,  phospho- 
molybdic  acid,  picric  acid,  trichloracetic  acid,  potassium  mercuric  iodide, 
and  tannic  acid.  The  protein  also  forms  insoluble  albuminates  with  the 
salts  of  the  heavy  metals. 

The  color  reactions  for  the  proteins,  which  are  given  in  the  laboratory 
experiments  at  the  end  of  this  chapter  are  due  to  a  reaction  between  some 
one  or  more  of  the  constituent  radicals  of  the  complex  protein  molecule  and 
the  chemical  reagent  or  reagents  used.  Thus  certain  color  tests  are  due 
to  the  presence  of  individual  amino  acids  in  the  protein  molecule,  and  the 
intensity  of  the  reaction  obtained  will  vary  with  the  amount  of  the  amino 
acids  present.  The  negative  results  for  a  certain  test  will  indicate  the  ab- 
sence of  the  particular  amino  acid  in  the  molecular  complex.  The  color 
tests,  then,  are  important  because  they  throw  light  on  the  chemical  consti- 
tution of  the  protein  under  observation. 

CLASSIFICATION  OF  THE  PROTEINS. 

The  following  classification  is  that  adopted  by  the  American  Physio- 
logical Society  and  the  American  Society  of  Biological  Chemists: 

I.  Simple  Proteins. — Protein  substances  which  yield  only  a-amino  acids 
or  other  derivatives  on  hydrolysis. 

a.  Albumins. — Soluble  in  pure  water;  e.g.,  ovalbumin,  seralbumin,  and 
the  vegetable  albumins. 

b.  Globulins. — Insoluble  in  pure  water,  but  soluble  in  neutral  solutions 
of  strong  bases  with  strong  acids;  e.g.,  ovoglobulin,  edestin,  and  other  vege- 
table globulins. 

c.  Glutelins. — Simple  proteins  insoluble  in  neutral  solvents,  but  readily 
soluble  in  very  dilute  acids  and  alkalies.     These  substances  occur  abundantly 
in  the  seeds  of  cereals. 


CLASSIFICATION    OF    THE    PROTEINS  87 

All  of  the  above  are  coagulable  by  heat. 

d.  Prolamins  or  Alcohol- soluble  Proteins. — Soluble  in  70  to  80  per  cent, 
alcohol;  insoluble  in  water,  absolute  alcohol,  and  other  neutral  solvents;  e.g., 
zein  from  corn,  gliadin  from  wheat,  and  hordein  from  barley. 

e.  Albuminoids. — Simple  proteins  characterized  by  a  pronounced  in- 
solubility in  all  neutral  solvents.     These  form  the  principal  organic  con- 
stituents of  the  connective  tissues  of  animals  including  their  external  covering 
and  its  appendages.     Examples:  elastin,  collagen,  and  keratin. 

The  above  sub-classes  are  characterized  by  physical  rather  than  by 
chemical  differences.  When  the  protein,  for  instance,  is  termed  a  globulin 
it  means  that  it  is  a  typical  simple  protein  with  certain  characteristic  solu- 
bilities. Proteins  intermediate  in  character  between  albumins  and  globulins 
are  met  with,  and  the  use  of  these  terms  as  a  hard-and-fast  classification 
has  led  to  considerable  confusion. 

/.  Histones.  On  hydrolysis  these  yield  a  large  number  of  amino  acids, 
among  which  the  basic  ones  predominate.  The  histones  stand  chemically 
between  the  typical  simple  proteins  and  the  following  group  of  protamins. 
Examples  are:  globin,  thymus  histone,  scombrone. 

g.  Protamines. — Simpler  polypeptids  than  the  proteins  included  in  the 
preceding  groups.  They  yield  comparatively  few  amino  acids,  among  which 
the  basic  ones  predominate.  They  are  the  simplest  natural  proteins. 
Examples  are:  salmin,  sturine,  clupeine,  and  scombrine. 

II.  Conjugated  Proteins. — Substances  which  contain  the  protein  mole- 
cule united  to  some  other  molecule  or  molecules  otherwise  than  as  a  salt. 

a.  Nucleoproteins. — Compounds  of  one  or  more  protein  molecules  with 
nucleic  acid;  e.g.,  nucleohistone. 

b.  Glycoproteins. — Compounds  of  the  protein  molecule  with  a  substance 
or  substances  containing  a  carbohydrate  group  other  than  a  nucleic  acid; 
e.g.,  mucins  and  mucoids. 

c.  Phosphoproteins. — Compounds  of  the  protein  molecule  with   phos- 
phorous containing  substances  other  than  a  nucleic  acid  or  lecithin;  e.g., 
casein,  ovovitellin. 

d.  Hemoglobins. — Compounds  of  a  protein  molecule  with  hematin  or 
some   similar   substance.     These   include   the   respiratory  pigments;   e.g., 
hemoglobin  and  hemocyanin. 

e.  Lecithoproteins. — Compounds  of  the  protein  molecule  with  the  lipoid 
lecithin;  e.g.,  lecithans,  phosphatides. 

III.  Derived    Proteins.     Class   i.     Primary    Protein    Derivatives. — 
Derivatives  of  the  protein  molecule  apparently  formed  by  hydrolytic  changes 
which  involve  only  slight  alteration  of  the  protein  molecule. 

/.  Proteans. — Insoluble   products  which  apparently  result  from  the  in- 
cipient action  of  water,  from  dilute  acids  or  enzymes;  e.g.,  myosan,  edestan. 
g.  Metaproteins. — Products  of  the  further  action  of  acids  and  alkalies 


88  THE   CHEMICAL   COMPOSITION   OF    THE   BODY 

whereby  the  molecule  is  so  far  altered  as  to  form  products  soluble  in  very 
weak  acids  and  alkalies,  or  insoluble  in  neutral  solutions;  e.g.,  acid  meta- 
protein,  acid  albuminate,  alkali  metaprotein  or  alkali  albuminate. 

h.  Coagulation  Proteins. — Insoluble  products  resulting  from  i,  the  action 
of  heat  on  protein  in  solution  or,  2,  the  action  of  alcohol  on  the  protein. 

Class  2.  Secondary  Protein  Derivatives. — Products  of  more  extensive 
hydrolytic  cleavage  of  the  protein  molecule  than  that  in  the  preceding  class. 

i.  Proteoses. — Soluble  in  water,  non-coagulable  by  heat,  and  precipitated 
by  saturating  their  solutions  with  ammonium  or  zinc  sulphate. 

;.  Peptones. — Soluble  in  water,  non-coagulable  by  heat,  and  not  pre- 
cipitated by  saturating  their  solutions  with  ammonium  sulphate. 

k.  Peptides. — Definitely  characterized  combinations  of  two  or  more 
amino  acids,  the  carboxyl  group  of  one  being  united  with  the  amino  group 
of  the  other  with  the  elimination  of  a  molecule  of  water. 


CHARACTERISTICS  OF  THE  VARIOUS  CLASSES  OF  PROTEINS. 

Albumins. — Albumins  constitute  the  first  class  of  simple  proteins. 
They  may  be  defined  as  simple  proteins  which  are  coagulable  by  heat  and 
are  soluble  in  pure  (salt-free)  water.  As  a  rule,  they  are  not  precipitated  on 
saturating  their  solutions  with  sodium  chloride  or  magnesium  sulphate, 
unless  the  solution  be  then  acidified  with  dilute  acid.  They  do  not  coagu- 
late out  on  heating  their  solution  unless  a  trace  of  a  salt  is  present.  Some 
albumins  have  been  prepared  in  crystalline  form;  e.g.  ovalbumin,  serum- 
albumin,  and  lactalbumin.  Crystallization  is  obtained  on  adding  ammonium 
sulphate  to  the  protein  solution  until  slight  turbidity  results,  then  clearing 
the  solution  by  adding  a  little  water  and  acidifying  slightly  with  acetic  acid. 
The  albumins,  as  a  rule,  are  precipitated  on  saturating  with  neutral  ammo- 
nium sulphate.  Zinc  sulphate  may  be  employed  when  it  is  desired  not  to 
introduce  ammonium  salts  into  the  precipitation.  The  proteins  are  pre- 
cipitated and  subsequently  coagulated  by  the  addition  of  an  excess  of  alco- 
hol. They  remain  in  solution  on  removal  of  inorganic  salts  by  dialysis. 

On  heating  a  solution  of  an  albumin  (or  a  globulin)  the  turbidity  and 
flocking  out  of  the  coagulum  occur  at  a  temperature  which  is  more  or  less 
characteristic  for  the  individual  protein.  However,  the  coagulation  tem- 
perature can  be  varied  according  to  the  concentration  of  the  protein  solu- 
tion, the  presence  of  inorganic  salts,  and  by  the  reaction  of  the  solution.  The 
coagulation  temperatures,  then,  cannot  be  given  as  definite  characters  for 
individual  proteins  unless  the  conditions  under  which  the  figures  were  ob- 
tained are  comparable. 

The  albumins  differ  among  themselves  in  the  cleavage  products  they 
yield  on  hydrolysis,  in  their  elementary  composition,  and  in  the  specific 
rotation  and  coagulation  temperatures.  The  serum  albumin  and  lact- 


ALBUMINOIDS  89 

albumin  are  quite  closely  related  chemically,  though  differing  in  their  specific 
rotation  of  polarized  light.  The  albumins  contain,  as  a  rule,  more  sulphur 
than  do  the  other  classes  of  proteins. 

Globulins. — The  globulins  are  simple  proteins  which  are  insoluble  in 
pure  (salt-free)  water,  but  which  are  soluble  in  neutral  solutions  of  salts  of 
strong  bases  with  strong  acids.  Most  globulins  are  precipitated  from 
their  solutions  on  slight  acidification  and  on  saturation  with  sodium  chloride 
and  magnesium  sulphate.  They  are  precipitated  also  from  other  solutions 
on  adding  equal  volume  of  saturated  ammonium  sulphate  solution;  this 
precipitation  is  commonly  termed  "  precipitation  at  half  saturation  ammon- 
ium sulphate."  Since  they  are  insoluble  in  pure  water,  dilution  of  the 
weak  salt  solution  containing  protein  causes  precipitation.  The  globulins 
are  especially  predominant  in  the  vegetable  kingdom.  They  occur  in  rela- 
tively large  amounts  as  the  reserve  protein  in  seeds  of  various  sorts.  There 
are,  however,  no  essential  differences  as  a  class  between  the  globulins  of 
animal  and  of  vegetable  origin. 

The  globulins  are  precipitated  from  weak  salt  solutions  on  dialysis  in 
pure  water.  The  inorganic  salts  diffuse  through  parchment,  and  with  the 
reduction  in  salt  content  the  globulins  are  precipitated.  Many  of  the 
vegetable  globulins  can  be  obtained  in  crystalline  form  by  precipitating 
them  in  this  way. 

As  a  class  the  globulins  are  relatively  less  stable  than  the  albumins. 
They  are  converted  over  into  proteans  on  successive  reprecipitation  or 
simply  by  standing  under  water. 

Albuminoids. — The  albuminoids  yield  similar  amino  acids  on  hydrol- 
ysis to  those  obtained  from  the  simple  proteins.  They  possess  essentially 
the  same  general  chemical  structure.  They  differ  from  all  other  proteins  in 
that  they  are  insoluble  in  neutral  solvents.  The  classification,  then,  is  based 
purely  on  this  property,  though  they  are  characterized  by  their  occurrence  as 
the  principal  organic  constituents  in  the  structure  of  the  supporting  tissues 
of  the  body  and  of  the  skin  and  its  appendages.  The  individual  albumin- 
oids differ  from  each  other  fundamentally  in  certain  chemical  character- 
istics. The  albuminoids  also  are  differentiated  along  with  the  morphologi- 
cal variations  of  the  connected  tissues  in  which  they  occur.  For  example, 
the  keratins  occur  in  the  skin  and  its  appendages.  Collagen  is  the  principal 
albuminoid  of  white  fibrous  tissue,  though  found  also  in  cartilage  and  bone. 
Elastin  characterizes  the  yellow  elastin  tissue,  as,  for  instance,  the  nuchal 
tendon,  the  elastic  tendon  so  well  developed  in  the  neck  of  the  ox. 

Keratins. — The  epidermis  of  the  skin,  the  nails,  hair  and  horn,  feathers, 
tortoise  shell,  silk,  and  the  supporting  neuroglia  of  nervous  tissue  may  be 
considered  to  be  keratins  in  relatively  pure  form.  The  keratins  take  the 
form  of  the  tissue  from  which  they  are  prepared.  On  heating  they  are 
decomposed  with  the  odor  of  burnt  horn.  They  are  insoluble  in  water, 


go  THE   CHEMICAL   COMPOSITION   OF   THE   BODY 

alcohol,  and  ether,  and  in  the  ordinary  protein  solvents.  They  are  not 
acted  upon  by  gastric  or  pancreatic  juices.  On  heating  to  150  to  200°  C. 
in  water  the  protein  is  hydrolyzed  and  dissolves.  The  keratins  are  also 
soluble  in  the  caustic  alkalies,  especially  on  heating.  Keratins  from  any 
source  may  be  prepared  in  pure  form  by  treating  with  artificial  gastric  juice, 
artificial  pancreatic  juice,  boiling  alcohol,  and  boiling  ether,  from  twenty- 
four  to  forty-eight  hours  being  devoted  to  each  process.  Several  keratins, 
so  far  as  their  chemical  structure  is  concerned,  exist. 

Collagen. — Collagen  can  be  most  satisfactorily  prepared  from  the  tendo 
achillis  of  the  ox.  It  forms  the  principal  organic  constituent  of  this  and 
other  white  fibrous  tissues,  as  shown  by  the  analysis  given  in  the  following 
table: 


CHEMICAL   COMPOSITION    OF   THE    WHITE   FIBROUS   TISSUE:     Tendo  Achillis 

(BERG  AND  GIES). 

Water 62  . 87  per  cent. 

Solids 37  . 13  per  cent. 

Inorganic  matter 0.47 

Organic  matter 36 . 66 

Fatty  substance  (ether-solu- 
ble)      i .  04 

Coagulable  protein 0.22 

Mucoid 1.28 

Elastin 1.63 

Collagen 31 . 59 

Extractives,  etc 0.90 

The  collagen  from  various  sources  in  common  with  the  keratin  is  not 
identical  in  composition.  It  differs  from  the  keratin  in  containing  less 
sulphur.  It  does  not  give  the  reaction  for  tryptophane  and  contains  but 
little  tyrosin.  It  is  dissolved  by  pepsin  and  hydrochloric  acid,  but  not  by 
pancreatic  juice. 

The  general  characteristic  of  collagen  is  that  it  is  hydrolyzed  into  gelatin 
by  boiling  with  water  or  dilute  acid.  Gies  has  shown  that  ammonia  is 
liberated  by  this  procedure.  Gelatin  is  soluble  in  hot  water,  but  its  solutions 
form  a  jell  when  cooled.  Inasmuch  as  tyrosin  and  tryptophane  are  not 
present  in  the  gelatin  molecule,  this  albuminoid  is  not  a  satisfactory  substi- 
tute for  the  protein  constituents  in  the  normal  diet. 

Elastin. — Elastin  is  the  principal  solid  constituent  of  yellow  elastic 
tissue;  e.g.,  the  ligamentum  nucha.  It  gives  the  ordinary  protein  color  re- 
actions. It  contains,  however,  a  relatively  small  amount  of  sulphur.  Elas- 
tin is  dissolved  by  pepsin  hydrochloric  acid  and  by  pancreatic  juice,  and 
unlike  collagen  it  is  not  converted  into  gelatin  on  prolonged  boiling  with 
water  or  dilute  acids. 


PROTAMINES  QI 

COMPOSITION  OF  YELLOW  ELASTIC  TISSUE:    Ligamentum  nuchce    (BERG  AND 

GIES). 

Water 57-57  Per  cent. 

Solids 42.43  per  cent. 

Inorganic  matter 0.47 

Organic  matter 41.96 

Fatty  substance  (ether-solu- 
ble)      1. 12 

Coagulable  protein 0.62 

Mucoid 0.53 

Elastin 31.67 

Collagen 7  .  23 

Extractives,  etc 0.80 

Histones. — The  histones  are  proteins  which  stand  in  their  chemical 
structure  between  the  true  proteins  and  the  protamines.  On  hydrolysis 
they  yield  a  large  number  of  amino  acids,  among  which  the  basic  ones  pre- 
dominate. The  basicity,  however,  is  only  slight.  They  are  precipitated 
by  dilute  ammonia  and  this  precipitate  is  soluble  in  an  excess  of  ammonia 
in  the  absence  of  ammonium  salts.  They  are  precipitated  by  nitric  acid, 
the  precipitate  dissolving  on  heating  and  again  appearing  on  cooling.  They 
give  precipitates  with  solutions  of  other  proteins.  On  heating,  the  histones 
yield  a  coagulum  which  is  easily  soluble  in  very  dilute  acids 

The  histones  are  found  in  the  nuclei  of  the  red  blood  cells  of  birds,  the 
unripe  testis  in  salmon  and  mackerel  and  in  the  ripe  spermatozoon  of  the 
sea-urchin.  The  thymus  contains  histone.  The  liver,  kidneys,  ox  pan- 
creas, and  testis  of  mammals  contain  no  histone-like  substances.  The 
globin  of  the  compound  protein  hemoglobin  is  to  be  regarded  as  a  histone. 

Protamines. — The  protamines  are  basic  proteins  which  are  combined 
with  nucleic  acid  and  form  the  chief  constituent  of  the  spermatozoa  of  the 
salmon  and  other  fish.  They  are  relatively  simple  proteins  yielding  com- 
paratively few  amino  acids  on  hydrolysis,  among  which  the  basic  ones 
predominate.  From  elementary  analyses  the  following  formula  has  been 
suggested  for  the  platinum  salt  of  salmine.  C30H57N17O6.4HC1.2PtCl4. 
As  seen  from  the  formula,  the  protamines  are  extremely  rich  in  nitrogen 
which  comprises  from  25  to  32  per  cent,  of  their  weight.  The  protamines 
dissolve  in  water  and  give  a  faintly  alkaline  reaction.  They  are  precipitated 
in  acid  solution  by  platinic  chloride.  The  protamines,  after  the  addition  of 
ammonia,  precipitate  true  proteins,  proteoses,  and  nucleic  acid.  On  hy- 
drolysis all  protamines  yield  relatively  large  amounts  of  argenine  and  varying 
amounts  of  histone,  lysin,  proline,  alanine,  valine,  leucin,  tyrosin,  and  ap- 
parently also  tryptophane.  Protamines  also  do  not  contain  sulphur  or  a 
carbohydrate  moiety.  They  are  not  changed  by  digestion  with  pepsin 
hydrochloric  acid. 


92  THE    CHEMICAL   COMPOSITION    OF    THE    BODY 

Conjugated  proteins  consist  of  a  protein  molecule  united  with  some 
other  molecule  or  molecules  otherwise  than  as  a  salt.  There  are  nucleo- 
proteins,  glycoproteins,  phosphoproteins,  hemoglobins  or  chromoproteins  and 
lecithoproteins — five  classes  of  conjugated  proteins. 

Nucleoproteins. — Nucleoproteins  contain  phosphorus  and  in  most 
instances  iron.  They  are  combinations  of  simple  proteins  with  a  substance 
known  as  nucleic  acid.  On  boiling  with  strong  acids  they  undergo  hydroly- 
tic  cleavage,  yielding  the  ordinary  protein  cleavage  products  from  the  pro- 
tein in  the  combination,  and  purine  and  pyrimidine  bases,  carbohydrates 
and  phosphoric  acid  from  the  nucleic  acid  moiety.  Nucleoproteins  are 
differentiated  from  the  phosphoproteins  in  that  the  latter  do  not  yield  purine 
and  pyrimidine  bases  on  hydrolytic  cleavage.  Nucleoproteins  are  the  es- 
sential organic  constituents  of  cell  nuclei.  They  go  into  solution  when  the 
tissues  are  extracted  with  cold  water  or  dilute  salt  solution.  They  are  pre- 
cipitated from  these  extracts  by  careful  acidification  and  dissolved  if  an 
excess  of  acid  is  added.  The  solutions  of  nucleoproteins  coagulate  on 
heating.  They  give  the  color  reactions  of  proteins. 

By  boiling  nucleoproteins  with  water  or  very  dilute  acetic  acid,  some 
of  the  protein  is  split  off.  There  result  substances  which  are  precipitated 
by  very  dilute  acids.  These  bodies  are  known  as  /^-nucleoproteins  and 
have  a  smaller  carbon  and  higher  phosphorus  content  than  the  original  nu- 
cleoprotein.  On  digestion  of  the  original  nucleoprotein  or  of  the  /?-nucleo- 
protein  with  pepsin  hydrochloric  acid,  a  precipitate  of  nuclein  is  obtained. 
On  further  digestion  with  pepsin  hydrochloric  acid,  or  with  trypsin,  or  on 
cleavage  with  acids  and  alkalies,  there  is  a  complete  splitting  away  of  the 
protein,  and  substances  are  formed  known  as  nucleic  acids.  The  structure 
and  cleavage  of  the  nucleoprotein  is  indicated  in  the  following  diagram. 

Nucleoprotein 

on  boiling  with  water  or  on  digestion 
with  pepsin  hydrochloric  acid  yields 


Nuclein  Protein 

On  long  boiling  with  water,  i 
per  cent,  hydrochloric  acid,  or 
dilute  alkalies  gives 


Nucleic  acid  Protein 

Nucleic  acids  are  not  merely  known  as  cleavage  products  of  nucleopro- 
tein, but  occur  preformed  in  the  cell  nuclei.  A  special  group  of  nucleic 
acids  are  bound  with  protamines  to  form  the  principal  constituent  of  the 
spermatozoa. 


PURINES  93 

Nucleic  acids  are  white,  amorphous  substances  containing  9  to  10  per  cent,  of 
phosphorus.  According  to  Levine  the  composition  corresponds  to  C43H67Ni6P4Ojo. 
Nucleic  acids  give  none  of  the  protein  reactions.  In  the  sperm  they  are  united  with  the 
strongly  basic  protamines  and  so  are  acid  in  character.  They  may  be  precipitated, 
however,  by  tannic  acid,  picric  acid,  or  phosphotungstic  acid  as  are  other  organic  bases. 
Nucleic  acid  has  been  isolated  from  many  different  tissues  and  apparently  is  a 
uniform  constituent  of  the  nuclei  of  all  cells.  The  nucleic  acids  from  different  animal 
tissues  are  apparently  identical  and  similarly  the  nucleic  acids  of  vegetable  origin  are 
identical,  though  there  are  marked  chemical  differences  between  these  two  types. 

The  structure  of  plant  nucleic  acid  is  the  better  known  of  the  two.  Thanks  to  the 
extensive  investigations  of  Levine,  and  Jones,  and  their  co-workers,  plant  nucleic  acid 
has  been  shown  to  be  composed  of  four  different  nucleotides.  These  nucleotides  are 
composed  of  a  purine  or  pyrimidine  base  linked  to  phosphoric  acid  by  carbohydrate 
groups.  In  the  case  of  yeast  or  plant  nucleic  acid,  the  carbohydrate  is  a  pentose 
(d.  Ribose).  The  first  of  the  nucleotides  studied  was  guanylic  acid  which  contains  the 
purine  base  quanine  or  2  ammo-  6  oxypurine. 

O  =  C— NH 
O  H   H  H    H   H  || 

MM!        /NH-C     C  =  NH 
HO— P— O— C— C— C— C— C— C 

\N—   C— NH 
OH        H   O  OH  OH 


Guanylic  acid. 

The  other  three  nucleotides  contain  adenine  and  the  pyramidine  bases  uracil  and 
cytosine  respectively.     The  tentative  structure  of  yeast  nucleic  acid  is  probably  that  of 
a  tetra  nucleotide,  though  the  mode  of  linkage  of  the  different  groups  is  still  in  doubt. 
HO\ 

HO/  guanine. 

H0\ 

HO/  cytosine. 

HO\ 

O  =  PO.C5H8O3.C4H3NaO2 
HO/  uracil. 

HO\ 

0=PO.C6H8O3— C6H4N6 
HO/  adenine. 

The  structure  of  animal  nucleic  acid  is  less  well  known.     It  contains  a  hexose  instead 
of  a  pentose  group  and  thymine  in  place  of  uracil. 

Purines. — To  the  purines  belong  a  number  of  extremely  important  animal 

and   plant   substances,   including    adenine,  guanine,   hypoxanthine,   xanthine, 

and  uric  acid,  and  the  methyl  purines  caffeine,  theobromine,  and  theophylline. 

The  mother  substance  of  the  purines  is  known  as  purine,  which  has  the 

following  structure: 

iN-C6  N  =  CH 

aC    C5-N\  HC    6-NH 

1  /°8  \CH 

6- N9/  /un 

N_C-N 
Purine  ring.  Purine. 


94 


THE    CHEMICAL    COMPOSITION    OF    THE   BODY 


Adenine  and  guanine  may  be  converted  into  hypoxanthine  and  xanthine, 
respectively,  when  added  to  extracts  of  tissues,  such  as  the  liver  and  the  thymus, 
spleen  and  pancreatic  glands.  The  "deamidization"  is  brought  about  by 
specific  enzymes  or  non-living  ferments  which  have  been  termed  adenase  and 
guanase.  On  slight  oxidation,  hypoxanthine  is  converted  into  xanthine,  and 
the  latter  into  uric  acid.  This  change  can  also  be  brought  about  in  the  body 
by  oxidizing  enzymes.  The  relation  of  the  purine  bases  to  uric  acid  is  indicated 
in  the  following  scheme: 


Adenine. 

(6-amino  purine.) 
N  =  C-NH2 

+  H20 

HC     C-NH 

-NH3 


Hypoxanthine. 
(6-oxypurine.) 
HN-C=O 


HC     C-NH 


J.L 


HN-C=0 


-c-/ 


CH 


on  oxi- 


dation 


HN-C=0 


HN-C=0 


H2N-C    C- 


+  H20 


NH 


I      I       >« 
N-C-N 


-NH3 


0  =  C 


Guanine. 
(2-amino  6-oxypurine.) 


on  oxidation 
:-NH  »     O 

>CH 
HN-C-N 

Xanthine. 
(  2-6di-oxypurine.) 


-NH 

>C  =  0 
HN-C-NH 

Uric  acid. 
(  2-6-8-trioxy  purine.) 

Glycoproteins. — The  glycoproteins  are  to  be  considered  as  compounds 
of  protein  and  considerable  quantities  of  a  carbohydrate  complex.  The  car- 
bohydrate group  can  be  split  from  the  protein  by  boiling  with  mineral  acids  or 
by  the  action  of  alkalies.  The  group  of  glycoproteins  includes  a  number  of 
proteins,  of  which  the  mucines  and  mucoids  are  the  most  important. 

Mucines  are  very  widely  distributed.  They  give  the  mucilaginous 
character  to  many  secretions  and  are  formed  and  discharged  through  the 
respiratory,  digestive,  and  other  tracts,  partly  by  mucous  cells  and  in  part  by 
the  mucous  glands,  especially  by  the  submaxillary  and  sublingual  sali- 
vary glands,  and  in  the  bile  passages.  On  hydrolysis  the  mucines  are  split, 
the  carbohydrate  moiety  yielding  glucose  amine  or  galactose  amine. 

Mucoids  occur  in  the  connecting  tissues  along  with  the  albuminoids. 
They  are  found  especially  in  the  tendon,  bone,  and  cartilage.  They  are 
combinations  of  protein  and  a  carbohydrate  containing  ethereal  sulphuric 
acid  known  as  chondroitin  sulphuric  acid.  On  cleavage,  besides  the  products 
formed  from  the  protein,  they  yield  sulphates  and  a  reducing  substance. 

Phosphoproteins.  —  The  phosphoproteins,  sometimes  called  nucleo- 
albumins,  are  compounds  of  the  protein  molecule  with  some  as  yet  unde- 
fined phosphorus-containing  substance  other  than  a  nucleic  acid  or  lecithin. 
While  the  phosphorus  content  of  these  substances  is  quite  similar  to  that 


METAPROTEINS  95 

of  the  nucleoproteins,  they  do  not  yield  any  purine  or  pyrimidine  bases  on 
hydrolytic  cleavage.  Two  of  the  best  known  phosphoproteins  are  the 
casein  of  milk,  and  vitellin  of  the  egg  yolk.  The  phosphorus  is  apparently 
present  as  a  phosphoric  acid  ester. 

Hemoglobins. — These  are  compounds  of  the  simple  protein  histone, 
with  an  iron-,  or  in  some  lower  animals,  copper-,  or  manganese-containing 
pigment  substance.  The  hemoglobins  are  more  fully  discussed  in  the  chapter 
on  the  Blood. 

Lecithoproteins. — These  are  combinations  of  proteins  and  a  fat-like 
substance,  lecithin.  Lecithin  is  a  compound  of  fatty  acids,  glycerin,  phos- 
phoric acid,  and  an  ammonium-like  organic  base,  choline.  The  combination 
of  lecithin  and  protein  is  apparently  a  loose  one:  the  lecithin  ordinarily  can 
be  split  off  by  boiling  alcohol.  The  lecithoproteins  include  substances 
commonly  termed  lecithans  and  phosphatids. 

The  derived  proteins  are  formed  as  intermediate  products  in  the  hydro- 
lytic cleavage  of  the  original  protein  molecule.  The  primary  protein  de- 
rivatives are  "apparently  formed  through  hydrolytic  changes  which  involve 
only  slight  alteration  of  the  protein  molecule." 

Metaproteins. — These  are  formed  from  the  simple  proteins  by  the 
action  of  weak  acids  and  alkalies.  This  class  comprises  what  have  com- 
monly been  termed  acid  and  alkali  albuminates.  The  metaproteins  are 
soluble  in  acid  or  alkaline  solution,  but  are  insoluble  in  neutral  solutions.  In 
the  formation  of  alkali  metaproteins,  the  sulphur  in  organic  combination 
is  split  off.  Thus  the  alkali  metaprotein  differs  from  the  acid  metaprotein 
in  that  the  former  contains  little  or  no  sulphur.  It  is  impossible,  then,  to 
transform  an  alkali  metaprotein  into  an  acid  metaprotein,  though  the  acid 
metaprotein  can  be  changed  into  the  other  modification.  Acid  metaproteins 
are  the  first  products  formed  in  the  pepsin  hydrochloric  acid  digestion  of 
proteins. 

Coagulated  Proteins. — Unaltered  typical  simple  proteins  in  solution  are 
altered  when  heated  or  by  long  standing  under  alcohol.  They  are  transformed 
into  a  coagulated  modification  no  longer  soluble  in  water  or  dilute  salt  solu- 
tions. A  similar  change  occurs  when  solutions  of  the  proteins  are  con- 
tinuously shaken  or  by  the  action  of  enzymes,  as  in  the  formation  of  fibrin 
from  fibrinogen  in  the  clotting  of  the  blood.  On  treating  coagulated  proteins 
with  acids  or  alkalies  they  are  converted  into  the  respective  metaproteins. 

Secondary  Protein  Derivatives. — Secondary  protein  derivatives  are 
intermediary  cleavage  products  which  result  from  a  more  profound  change 
than  occurs  in  the  formation  of  the  primary  derived  protein. 

Proteases  or  albumoses  are  intermediate  products  in  the  digestion  of 
proteins  by  proteolytic  enzymes  or  in  the  cleavage  with  acids.  Peptones 
are  yet  more  simple  products  than  the  proteoses  and  are  to  be  regarded 
as  relatively  simple  polypeptides  which  still  retain  some  of  the  protein 


96          THE  CHEMICAL  COMPOSITION  OF  THE  BODY 

characteristics.  A  number  of  proteoses  and  peptones  have  been  described. 
However,  there  is  no  sharp  dividing  line  between  the  more  simple  proteoses 
and  more  complex  peptones,  or  between  the  simple  peptones  and  the  peptides. 
(The  term  peptide  as  at  present  understood  designates  only  those  combina- 
tions of  amino  acids  possessing  a  known  definite  structure.)  The  peptones 
differ  from  the  proteoses  in  being  more  diffusible,  being  non-precipitable 
on  saturation  with  ammonium  sulphate,  and  by  their  failure  to  give  certain 
protein  reactions.  As  a  class,  proteoses  and  peptones  are  relatively  very 
soluble  and  are  non-coagulable  by  heat. 

Melanins  are  the  pigmentary  substances  found  in  the  hair,  feathers,  skin, 
the  choroid  coat  of  the  eye,  and  in  some  tumors.  Products  similar  to  the 
naturally  occurring  melanins  are  obtained  on  hydrolizing  nearly  all  proteins 
with  acids.  The  melanins  are  sulphur- containing  acid-like  substances,  and 
seem  to  be  combinations  of  amino-sugars  (glucosamine)  with  certain  amino- 
acids,  especially  tyrosine,  tryptophane,  and  lysine.  Iron  is  found  in  some  of 
the  melanins. 

THE  FATS. 

Fats  occur  very  widely  distributed  in  the  plant  and  animal  kingdom,  and 
constitute  one  of  the  four  classes  of  food  stuffs.  Fats  are  esters  or  ethereal 
salts  consisting  of  an  organic  radical  (glycerol)  united  with  the  residue  of  an 
organic  acid.  Ethyl  alcohol  may  be  combined  as  an  ester  with  acetic  acid. 

CH3COOH  +  C2H5OH  =  CH3COOC2H5  +  H2O 

Acetic  acid  Alcohol  Etyhl  acetate 

Similarly  the  triatomic  alcohol  glycerol  may  be  combined  with  the 
higher  fatty  acids  to  form  the  true  fats. 

CH2OH  CH2-OOCC15H31 

CHOH  +  3C15H31COOH  =  CH-  OOCC15H31  +  3H2O 

CH2OH  CH2-OOCCi5H31 

Glycerol   Palmitic  acid  Tri-palmitin 

The  animal  fats  are  for  the  most  part  mixtures  of  tri-palmitin,  tri- stearin, 
and  tri-olein,  the  last  two  being  esters  of  glycerol  with  stearic  acid,  C17H35 
COOH,  and,  with  the  unsaturated  oelic  acid,  C17H33COOH.  Human  fat 
consists  of  a  mixture  of  which  tri-palmitin  and  tri-stearin  comprise  three- 
fourths  of  the  whole.  The  fat  in  milk  and  butter  is  in  part  tri-butyrin,  the 
ester  of  glycerol  with  butyric  acid,  C2H5COOH.  The  percentage  of  any 
individual  fat  in  animal  tissue  depends  on,  and  is  characteristic  of,  the  particu- 
lar species  of  animal  from  which  the  fat  was  obtained.  Ordinary  mutton  fat 
contains  more  tri-stearin  and  less  tri-olein  than  pork  fat,  and  the  mutton 
fat  is  stiffer  because  the  melting-point  of  the  tri-stearin  is  the  highest  of 
the  fats. 

The    pure  fats  are  odorless,  tasteless,  and  generally  colorless.     They 


THE    FATS  97 

are  insoluble  in  water  and  cold  alcohol,  but  are  dissolved  by  acetone,  hot 
alcohol,  benzol,  chloroform,  and  ether.  When  shaken  with  water,  protein 
solutions,  soap,  or  gum  arabic,  the  fats  assume  a  finely  divided  condition 
known  as  an  emulsion.  The  suspension  in  water  is  only  temporary,  while 
the  emulsions  are  permanent. 

The  fats  are  hydrolyzed  or  saponified  by  superheated  steam  into  glycerol 
and  the  fatty  acids,  the  reaction  being  the  reverse  of  that  indicated  in  the 
equation  above.  On  boiling  with  caustic  alkalies,  they  are  similarly  saponi- 
fied; the  fatty  acids  are  then  combined  with  the  bases  to  form  salts  or  soaps. 

Lecithins  are  tri-glycerides  in  which  the  H  atom  of  two  instead  of  three 
groups  of  the  glycerol  is  replaced  by  a  fatty  acid  radical;  for  the  H  of  the 
third  hydroxyl  (OH)  group  there  is  substituted  an  ester-like  combination 
of  phosphoric  acid  with  a  nitrogen-containing  organic  base,  choline. 


CH2OOCC17H 


35 


CHOOCC17H35  C2H4OH 

CH2O  -  O  P  -  O  C2H4  N=(CH3)3 

\  \ 

(CH3)3=N  OH 

HO 

Lecithin  Choline 

On  saponification  the  di-stearyl  lecithin  molecule  above  combines  with 
three  molecules  of  water  and  is  split  into  two  molecules  of  stearic  acid,  one 
of  glycero-phosphoric  acid  and  one  of  choline. 

The  lecithins  are  soluble  in  alcohol,  benzene,  chloroform,  and  ether. 
They  are  precipitated  from  chloroform  or  alcohol-ether  solution  by 
acetone. 

The  lecithins  are  found  in  nearly  all  animal  and  vegetable  tissues,  espe- 
cially in  nervous  tissues.  They  are  essential  constituents  of  the  cell. 
Kephalin  is  of  more  than  passing  interest  in  that  its  presence  hastens 
blood  clotting.  It  accomplishes  this  result  by  removing  the  restraints 
of  antithrombin  on  fibrin  formation. 

Cholesterol  is  a  complex  alcohol  with  the  elementary  formula  C27H45OH, 
and  related  to  the  vegetable  terpenes,  being  grouped  with  the  fats  solely 
because  of  its  physical  properties.  Accordingly,  it  cannot  be  saponified.  It 
crystallizes  in  the  form  of  thin,  colorless,  transparent  plates  usually  notched 
in  one  corner.  It  exists  in  the  tissues  in  part  in  the  form  of  esters  with  the 
complex  fatty  acids.  Cholesterol  is  an  essential  cell  constituent;  it  is  present 
in  relatively  large  amounts  in  nervous  tissue.  It  occurs  also  in  wool  fat,  eggs, 
milk,  and  blood  plasma. 

Cholesterol  and  the  lecithins  are  often  termed  lipoids  or  fat-like 
substances. 


98  THE    CHEMICAL    COMPOSITION    OF    THE    BODY 


CARBOHYDRATES. 

The  typical  carbohydrates  contain  carbon  combined  with  hydrogen  and 
oxygen  in  the  proportion  to  form  water.  Other  substances,  such  as  acetic 
acid,  CH3COOH,  lactic  acid,  CH3CHOHCOOH,  and  inosit,  (CHOH)6, 
which  contain  hydrogen  and  oxygen  in  the  proportion  to  form  water,  are  not 
carbohydrates.  Certain  true  carbohydrates  also  do  not  fulfill  this  condition. 
Chemically,  the  carbohydrates  are  aldehyde  or  ketone  derivatives  of  complex 
alcohols;  i.e.,  they  have  the  structure 

R  -  CHOH  -  CHO  or  R  -  CO  -  CH2OH 

Aldose  Ketose 

Accordingly,  the  carbohydrates  are  termed  aldoses  or  ketoses.  They 
are  commonly  classified  by  the  number  of  carbon  atoms  in  the  molecule; 
e.g.,  pentoses  are  those  containing  five  carbon  atoms,  and  hexoses  have  six 
carbons.  Each  member  of  the  carbohydrate  class,  with  the  exception  of 
the  pentoses,  may  be  regarded  as  containing  the  saccharide  group,  C6H10O5. 
The  monosaccharides  are  then  C6H10O5  +  H2O;  the  di-saccharides  contain 
two  saccharide  groups  with  water,  (C6H10O5)2 +H2O,  while  the  poly- 
saccharides  contain  this  group  taken  a  large  number  of  times,  (C6H10O5)n. 
In  general,  the  solubility  of  the  saccharides  varies  inversely  with  the 
number  of  saccharide  groups  present:  the  mono-saccharides,  as  a  class, 
being  the  most  soluble  and  the  poly-saccharides  being  the  least  so.  On 
boiling  in  the  autoclave  or  with  mineral  acids,  and  by  the  action  of  amylo- 
lytic  and  inverting  enzymes,  the  poly-saccharides  are,  as  a  rule,  hy- 
drolyzed  into  the  simple  carbohydrates.  The  reaction  may  be  indicated 

(C6H1005)n    +    nH20    =    nC6H1206 

Poly-saccharide          Water         Mono-saccharide 

Simple    poly-saccharides,  and   di-saccharides   are   formed  as  intermediate 
products  in  the  cleavage  and  in  turn  these  are  further  hydrolyzed. 

C12H220U    +   H20    ,  ,    2C6H12Oe 

Di-saccharide         Water       Mono-saccharide 

The  color  reactions  with  iodine,  fermentation  with  yeast  and  bacteria, 
the  formation  of  characteristic  crystalline  osazones  with  phenylhydrazine 
and  the  reducing  of  alkaline  solutions  of  the  oxides  of  metals  like  copper, 
bismuth,  mercury,  and  ammoniacal  silver  solutions,  are  the  most  widely 
used  reactions  for  testing  and  differentiating  the  carbohydrates.  The 
reduction  of  alkaline  solutions  of  the  metallic  oxides  is  due  to  the  easily 
oxidized  aldehyde  and  ketone  structure  of  the  sugar. 


CARBOHYDRATES  99 

The  more  common  carbohydrates  may  be  listed  as  follows: 

1.  Mono-saccharides. 

Hexoses,  C6H12O6 — dextrose,  levulose,  galactose. 

Pentoses,  C5H10O5 — arabinose,  xylose,  rhamnose  (methylpentose, 

C6H1205). 

2.  Di-saccharides,  C12H22On, 

Maltose,  saccharose  (cane-sugar),  lactose. 

3.  Poly-saccharides,  (C6H10O5)n. 

Dextrin  group — dextrins. 

Starch  group — starch,  inulin,  lichenin,  glycogen. 

Cellulose  group — cellulose. 

Dextrose  (glucose,  grape-sugar)  is  an  aldose  found  in  honey  and 
in  many  fruit  juices  where  it  is  usually  associated  with  levulose.  It  is 
present  in  the  blood  in  small  amounts,  o.i  to  0.15  per  cent.,  in  normal 
urine  in  minute  traces,  and  in  diabetic  urine.  It  is  not  as  sweet  as  cane- 
sugar.  Glucose  is  produced  on  boiling  starch  with  dilute  acids.  It  is  very 
soluble  in  water  and  is  slightly  soluble  in  alcohol.  It  crystallizes  from 
water  in  leaves  or  plates  and  from  alcohol  in  anhydrous  needles.  Dextrose 
rotates  the  plane  of  polarized  light  to  the  right — its  specific  rotation  is  given 
by  the  expression  [a]d  =  +52.5°.  It  forms  a  characteristic  glucosazone 
when  boiled  with  phenylhydrazine  in  the  presence  of  acetic  acid;  the  osazone 
crystallizes  from  the  hot  solution.  Glucose  reduces  metallic  oxides  in 
alkaline  solution.  It  undergoes  alcoholic  fermentation  with  yeast  and  acid 
fermentation  with  certain  bacteria. 

CH2OH  CH2OH 

CHOH  CHOH 

CHOH  CHOH 

CHOH  CHOH 

CHOH  CO 

CHO  CH2OH 

Dextrose.  Levulose. 

Levulose  (fructose)  is  a  ketose  and  is  found  associated  with  dextrose 
in  many  fruits,  the  mixture  probably  being  produced  by  the  hydrolysis  of,  or 
preceding  the  synthesis  of,  cane-sugar.  It  may  be  prepared  by  the  hydrolysis 
of  inulin  and,  along  with  dextrose,  by  the  inversion  of  cane-sugar  on  boiling 
with  dilute  mineral  acids  or  through  the  action  of  specific  enzymes.  It  is 


IOO  THE    CHEMICAL   COMPOSITION    OF   THE    BODY 

levorotatory,  [a]d  =  —92°.  Levulose  may  be  crystallized  with  difficulty  in 
needles.  It  has  a  sweet  taste.  It  reduces  alkaline  solutions  of  the 
metallic  oxides,  but  not  so  much  as  dextrose,  and  yields  an  osazone  identical 
with  glucosazone.  It  undergoes  fermentation,  but  less  readily  than  dextrose. 

Galactose  is  obtained  with  dextrose  from  milk-sugar  or  lactose  on 
boiling  with  dilute  mineral  acids.  It  is  less  soluble  in  water  than  levulose 
or  glucose.  It  reduces  metallic  oxide  in  alkaline  solution,  forms  a  charac- 
teristic osazone  which  melts  at  193-4°  C.  and  it  undergoes  slow  fermentation 
by  yeast.  It  is  dextrorotatory. 

It  is  also  obtained  on  hydrolysis  of  cerebrin,  a  glucoside  occurring  in 
nervous  tissue. 

Maltose  (malt -sugar)  is  produced  by  the  action  of  amylolytic  enzymes 
on  starch  and  glycogen.  It  crystallizes  in  small  needles,  is  strongly  dextro- 
rotatory, [a]d  =  +140.6°,  reduces  alkaline  copper  solutions  much  feebler 
than  dextrose,  and  does  not  reduce  bismuth  oxide.  It  forms  a  characteris- 
tically crystalline  osazone,  melting  at  206°  C.  It  is  readily  soluble  in  water 
and  only  slightly  soluble  in  alcohol.  Maltose  is  not  so  sweet  as  cane-sugar. 
It  does  not  undergo  alcoholic  fermentation  with  yeast  unless  first  split  into 
dextrose.  On  boiling  with  dilute  mineral  acids  or  through  the  action  of 
inverting  enzymes,  it  is  hydrolyzed  into  dextrose. 

Saccharose  (cane-sugar)  is  obtained  from  many  plants,  such  as  the 
sugar  beet  and  sugar  cane,  and  from  the  sap  of  certain  trees,  as  the  sugar 
maple.  It  crystallizes  in  prisms,  is  soluble  in  water,  and  only  very  slightly 
soluble  in  alcohol.  Cane-sugar  is  not  directly  fermentable  by  yeast.  It  is 
dextrorotary.  Saccharose  has  no  reducing  action  on  alkaline  copper  solution 
— the  dextrose  and  levulose  yielded  on  inversion  probably  being  united 
together  by  the  aldehyde  and  ketone  radicals,  respectively.  Similarly,  it  does 
not  form  an  osazone.  In  strong  solutions,  saccharose  acts  as  a  food  pre- 
servative against  bacterial  or  other  decomposition  through  organic  agencies. 
On  boiling  with  dilute  mineral  acids  or  through  the  action  of  inverting  en- 
zymes, one  molecule  of  saccharose  yields  one  molecule  of  dextrose  and  one 
molecule  of  levulose. 

Lactose  (milk-sugar)  is  present  as  the  chief  carbohydrate  of  milk.  It 
may  be  separated  from  the  whey — the  product  remaining  after  skimming 
milk  and  precipitating  the  proteins  therein.  It  can  be  prepared  in  large 
hard  crystals.  It  is  much  less  soluble  in  water  than  cane-sugar,  and  has  but 
a  slightly  sweet  taste.  Lactose  is  strongly  dextrorotatory,  [a]d  =  +52.5°. 
It  reduces  alkaline  copper  solution  and  forms  a  characteristic  osazone.  It 
is  not  fermented  by  ordinary  yeast;  certain  bacteria  easily  convert  it  into 
lactic  and  other  simple  organic  acids.  The  inverted  milk-sugar  undergoes 
alcoholic  fermentation  readily,  and  kumiss  and  kephir,  made  from  mare's 
and  cow's  milk,  respectively,  are  prepared  in  this  way.  On  hydrolyzing  by 
boiling  with  mineral  acids  or  through  the  action  of  inverting  enzymes,  the 
lactose  is  split  into  dextrose  and  galactose. 


CARBOHYDRATES 


IOI 


Dextrins  are  a  series  of  intermediate  poly-saccharides  between  the  di- 
saccharides  and  the  starches.  They  are  non-crystalline,  are  soluble  in 
water,  and  are  precipitated  on  the  addition  of  alcohol  in  excess.  They  are 
dextrorotatory  and  are  not  fermented  by  yeast.  Their  power  to  reduce 
alkaline  copper  solution  has  been  questioned,  but  they  yield  osazones  which 
are  relatively  soluble.  They  have  a  slightly  sweet  taste.  The  more  com- 
plex dextrins  give  a  red  reaction  with  iodine. 

Starch  is  found  in  various  parts  of  plants,  especially  in  the  tubers  and 
seeds.  It  is  a  form  of  storage  carbohydrate,  and  serves  as  a  source  of  ma- 
terial for  the  development  of  the  young  plant.  Starch  is  obtained  commer- 
cially from  potatoes,  rice,  corn,  wheat,  sago,  and  the  like.  It  constitutes  the 
greater  proportion  of  our  food. 

Starch  as  obtained  is  a  soft  white  powder  which  on  microscopic  examina- 
tion is  found  to  consist  of  small  granules.  These  are  often  characteristic  in 
shape  and  size  according  to  the  origin  of  the  material.  The  granules  appear 
to  be  built  up  of  concentric  layers  of  two  varieties  of  starch.  The  more  sol- 
uble form,  known  as  amylose,  is  ensheathed  by  the  less  soluble  variety  of 
starch,  the  amylopectin.  This  sheath  is  ruptured  by  boiling  the  aqueous  sus- 
pension of  starch  granules  and  an  opalescent  solution  or  starch  paste  is 
obtained.  On  standing  in  dilute  solution,  the  opalescent  material  settles  to 
the  bottom,  but  the  clear  fluid  above  still  gives  the  blue  reaction  with  iodine. 
This  color  is  characteristic  for  starch;  it  disappears  on  heating,  but  returns 
when  the  liquid  cools.  Starch  will  not  diffuse  through  a  semi-permeable 
membrane.  On  boiling  with  dilute  mineral  acids  starch  is  hydrolyzed  to  dex- 
trose. The  dextrins  and  maltose  are  formed  as  intermediate  products. 
With  amylolytic  enzymes,  the  change  practically  only  goes  as  far  as  the 
maltose  stage. 

Glycogen  (animal  starch)  is  the  reserve  form  of  carbohydrates  in 
animals.  It  is  synthesized  from  dextrose  and  can  again  be  hydrolyzed  to 
dextrose  for  transportation  or  for  oxidation  to  yield  energy  to  the  tissues. 
The  following  table  shows  the  per  cent,  and  distribution  of  glycogen  in  the 
various  tissues  of  the  dog  (Schondorff) : 


Organ 

Per  cent,  glycogen 

Per  cent,  of  total 
glycogen. 

Blood     

o  .  004  i 

o  .  01  <c 

Liver  .  . 

7207 

•77.07 

Muscles 

O    7  ^OO 

A  A   ,  2  "? 

Bones  
Viscera  

0.2736 

o  .  2024 

9-2$ 
^?.8l 

Skin       .  . 

o  .  08  so 

4.  40 

Heart 

O     2T.1 

O     17 

Brain 

o    1  084. 

O    OQ 

102  THE    CHEMICAL    COMPOSITION    OF    THE    BODY 

It  occurs  in  relatively  large  amounts  in  some  invertebrates,  especially 
in  moluscs  and  in  intestinal  worms.  The  muscles  and  reproductive  organs  of 
oysters,  clams,  and  scallops  are  very  rich  in  this  substance. 

Glycogen  resembles  starch  in  forming  opalescent  solutions.  It  may  be 
prepared  from  the  liver  or  muscle  of  a  freshly  killed  animal  by  boiling  the 
tissue  to  coagulate  the  proteins,  grinding  with  sand,  boiling  with  water 
slightly  acidified  with  acetic  acid  and  precipitating  the  nitrate  with  an  excess 
of  alcohol;  dilute  or  concentrated  solutions  of  caustic  alkalies  may  be  used 
to  extract  all  the  glycogen.  Glycogen  is  a  white,  tasteless,  and  amorphous 
powder.  It  gives  a  maroon  color  with  iodine,  and  does  not  reduce  alkaline 
copper  sulphate  solution.  It  is  completely  precipitated  by  saturating  its 
solution  with  solid  ammonium  sulphate,  by  tannic  acid,  or  by  ammoniacal 
basic  lead  acetate.  On  hydrolysis  with  mineral  acids  or  on  digestion  with 
amylolytic  enzymes  it  yields  the  same  series  of  products  as  ordinary  starch. 

Inulin  is  the  reserve  carbohydrate  of  the  Composites,  occurring  in  the 
tubers  of  the  artichoke  and  dahlia  and  in  the  roots  of  the  dandelion  and  bur- 
dock. On  hydrolysis  with  acids  or  the  enzyme  inulase,  it  yields  levulose. 
Inulin  is  slightly  soluble  in  cold  and  easily  soluble  in  hot  water.  It  is  precipi- 
tated from  its  solution  by  an  excess  of  alcohol.  The  digestive  enzymes  of  the 
body  do  not  act  on  inulin. 

Lichinin  is  obtained  from  the  Cetraria  Islandica  (Iceland  moss).  It  forms 
a  difficultly  soluble  jelly  in  cold  water  and  an  opalescent  solution  in  hot  water. 
On  hydrolysis  with  dilute  acids,  it  yields  dextrines  and  dextrose.  The 
ordinary  digestive  enzymes  have  no  action  on  lichinin. 

Cellulose  forms  a  large  portion  of  the  cell  wall  or  the  woody  structure 
of  plants.  It  is  extremely  insoluble.  Chemically,  it  is  more  complex  than 
the  common  starch  molecule.  The  hydrochloric  and  hydrofluoric  acid 
extracted  (ash-free)  filter-papers  and  absorbent  cotton  are  examples  of  prac- 
tically pure  cellulose. 

INORGANIC  SUBSTANCES  OF  THE  BODY. 

Salt. — The  inorganic  principles  of  the  human  body  are  numerous. 
They  are  derived,  for  the  most  part,  directly  from  food  and  drink  and  pass 
through  the  system  unaltered.  Some  radicals  are  newly  formed  by  oxi- 
dation within  the  body,  as,  for  example,  a  part  of  the  sulphates  and  car- 
bonates from  the  sulphur  of  the  proteins  and  the  carbon  of  protein,  fat,  and 
carbohydrate. 

Much  of  the  inorganic  saline  matter  found  in  the  body  is  a  necessary 
constituent  of  its  structure,  as  necessary  in  its  way  as  protein  or  any  other 
organic  principle.  Another  part  is  important  in  regulating  or  modifying 
various  physical  processes,  as  absorption,  solution,  and  the  like.  A  part 
must  be  reckoned  only  as  matter  which  is,  so  to  speak,  accidentally  present, 
whether  derived  from  the  food  or  the  tissues,  and  which  will,  at  the  first 


INORGANIC    SUBSTANCES    OF    THE    BODY  103 

opportunity,  be  excreted  from  the  body.     The  principal  salts  present  in 
the  body  are: 

Sodium  and  Potassium  Chlorides. — These  salts  are  present  in  nearly  all 
parts  of  the  body.  The  former  seems  to  be  especially  necessary,  judging 
from  the  instinctive  craving  for  it  on  the  part  of  animals  in  whose  food  it 
is  deficient,  and  from  the  condition  which  is  consequent  on  its  withdrawal. 
The  quantity  of  sodium  chloride  in  the  blood  is  greater  than  that  of  all  its 
other  saline  ingredients  taken  together,  but  it  is  present  chiefly  in  the  fluids 
of  the  body.  In  the  tissues,  the  muscles  for  example,  the  quantity  of  sodium 
chloride  is  less  than  that  of  the  chloride  of  potassium,  which  forms  a  constant 
ingredient  of  protoplasm. 

Calcium  Fluoride. — It  is  present  in  minute  amount  in  the  bones  and 
teeth,  and  traces  have  been  found  in  the  blood  and  some  other  fluids. 

Calcium,  Potassium,  Sodium,  and  Magnesium  Phosphates. — These  phos- 
phates are  found  in  nearly  every  tissue  and  fluid.  In  some  tissues — the  bones 
and  teeth — tricalcium  phosphate  exists  in  very  large  amount.  The  phos- 
phate of  calcium  is  intimately  incorporated  with  the  organic  basis  or  matrix, 
but  it  can  be  removed  by  acids  without  destroying  the  general  shape  of  the 
bone.  After  the  removal  of  its  inorganic  salts,  a  bone  is  left  soft,  tough, 
and  flexible. 

Potassium  and  sodium  phosphates,  with  the  carbonates,  maintain  the 
alkalinity  of  the  blood. 

Calcium  Carbonate. — It  occurs  in  bones  and  teeth,  but  in  much  smaller 
quantity  than  the  phosphate.  It  is  found  also  in  some  other  parts.  The 
small  concretions  of  the  internal  ear  of  some  fishes  (otoliths)  are  composed 
of  crystalline  calcium  carbonate,  and  form  the  only  example  of  inorganic 
crystalline  matter  existing  as  such  in  the  body. 

Potassium  and  Sodium  Carbonates  and  Sulphates. — These  are  found  in 
the  blood  and  most  of  the  secretions  and  tissues. 

Silicon. — A  very  minute  quantity  of  silica  exists  in  the  urine  and  in 
the  blood.  Traces  of  it  have  been  found  also  in  bones,  hair,  and  some 
other  parts. 

Iron. — The  especial  place  of  iron  is  in  hemoglobin,  the  coloring- matter 
of  the  blood,  of  which  a  full  account  will  be  given  with  the  chemistry  of  the 
blood.  Iron  is  found,  in  very  small  quantities,  in  the  ashes  of  bones,  mus- 
cles, and  many  tissues,  and  in  lymph  and  chyle,  albumin  of  serum,  fibrin, 
bile,  milk,  and  other  fluids. 

Iodine  occurs  as  an  iodized  protein  in  the  thyroid  gland.  Biologically, 
it  is  found  as  a  tri-iodotyrosin  in  sponges  and  the  Gorgonian  corals. 

Water. — Water  forms  a  large  proportion,  more  than  two-thirds,  of 
the  weight  of  the  whole  body.  Its  relative  amount  in  some  of  the  principal 
solids  and  fluids  of  the  body  is  shown  in  the  following  table  (from  Robin 
and  Verdeil) : 


104  THE    CHEMICAL    COMPOSITION    OF    THE   BODY 


Quantity  of  Water  in  Per  Cent. 

Teeth 10.0  Bile 88.0 

Bones 13.0  Milk 88.7 

Cartilage 55 -o  Pancreatic  juice. ...  90.0 

Muscles 75-°  Urine 93-6 

Ligament 76.8  Lymph 96.0 

Brain 78.9  Gastric  juice 97-5 

Blood 79 .  5  Perspiration 98.6 

Synovia 80 .  5  Saliva 99-5 

In  all  the  fluids  and  tissues  of  the  body — blood,  lymph,  muscle,  gland, 
etc. — water  acts  the  part  of  a  general  solvent,  and  by  its  means  alone  circula- 
tion of  nutrient  matter  is  possible.  It  is  the  medium  also  in  which  all  fluid 
and  solid  aliments  are  dissolved  before  absorption,  as  well  as  the  means  by 
which  all,  except  gaseous,  excretory  products  are  removed.  All  the  various 
processes  of  secretion,  transudation,  and  nutrition  depend  of  necessity  on 
its  presence  for  their  performance. 

The  greater  part,  by  far,  of  the  water  present  in  the  body  is  taken  into 
it  as  such  from  without,  in  the  food  and  drink.  A  small  amount,  however, 
is  the  result  of  the  chemical  union  of  hydrogen  with  oxygen  in  the  oxidations 
of  the  body. 

The  loss  of  water  from  the  body  is  intimately  connected  with  excretion 
from  the  lungs,  skin,  and  kidneys,  and,  to  a  less  extent,  from  the  alimentary 
canal.  The  loss  from  these  various  organs  may  be  thus  apportioned  (quoted 
by  Dalton  from  various  observers): 

From  the  alimentary  canal  (feces) 4  per  cent. 

From  the  lungs 20  per  cent. 

From  the  skin   (perspiration) 30  per  cent. 

From  the  kidneys  (urine) 46  per  cent. 

100 

Under  some  conditions  the  loss  of  water  from  the  alimentary  canal  may 
be  enormously  increased,  as  in  acute  diarrheas.  In  young  children  and 
babies  in  particular  this  fact  is  often  not  realized  and  not  enough  water  is 
given  by  the  mouth  to  supply  the  loss.  The  result  is  a  considerable 
concentration  of  the  blood  and  tissues,  a  relative  dessication  that  may 
prove  very  injurious. 


CHEMISTRY   OF    THE   BODY  105 

LABORATORY  EXPERIMENTS  ON  THE  CHEMISTRY  OF  THE 

BODY. 

This  list  will  serve  as  basal  for  the  guidance  of  students  and  teachers. 
The  experiments  listed  are  to  be  supplemented  by  technical  laboratory 
guides  and  references  to  fuller  discussion  in  the  literature. 

THE  PROTEINS. 

i.  Preparation  of  Proteins. — The  most  convenient  source  of  proteins 
for  laboratory  work  are  blood  serum,  egg  white,  or  commercial  preparations 
of  the  milk  protein  casein.  Protein  may  also  be  prepared  from  various 
plant  seeds,  especially  cereals.  Hempseed  contains  a  globulin,  edestin, 
which  is  very  easily  isolated  in  the  laboratory. 

a.  Preparation  of  Edestin. — Grind  up  some  hemp-seed  in  an  ordinary 
meat  chopper  and  extract  the  resulting  meal  with  5  per  cent,  salt  solution, 
warming  to  60°.     The  solution  should  not  be  heated  above  65°  because 
the  protein  will  be  coagulated.     Filter  while  hot.     On  cooling  slowly  the 
edestin  will  crystallize  out.     Examine  some  of  the  precipitate  with  a  micro- 
scope and  sketch  the  crystals.     The  edestin  is  soluble  in  10  per  cent,  salt 
solution  without  warming,  and  solutions  for  laboratory  use  can  be  prepared 
in  this  way. 

b.  Preparation  of  Egg  Albumin. — The  yolk  should  be  separated  from 
the  white  of  fresh  eggs  and  the  reticulum  in  the  egg  white  broken  up  with 
a  wire  egg-beater.     Egg  white  can  then  be  diluted  as  desired  and  the  pre- 
cipitate globulin  filtered  off.     Crystals  of  ovalbumin  can  be  prepared  as 
follows: 

The  egg  white,  beaten  as  directed,  is  strained  through  gauze  and  an 
equal  volume  of  saturated  ammonium  sulphate  solution  is  added.  After 
twenty-four  hours  the  globulin  precipitate  is  filtered  off  and  concentrated 
ammonium  sulphate  solution  added  until  the  mixture  becomes  turbid. 
Then  distilled  water  is  added  very  carefully  until  turbidity  has  disappeared. 
The  solution  is  then  acidified  with  acetic  acid,  which  has  been  saturated 
with  ammonium  sulphate,  until  a  precipitate  is  obtained."  The  precipitate 
is  at  first  amorphous,  and  on  standing  becomes  crystalline.  Examine  the 
crystals  under  the  microscope  and  sketch  them. 

c.  Other   Protein   Crystals. — Crystals  of    hemoglobin    may  be  demon- 
strated by  adding  a  drop  of  ether  to  diluted  dog  blood  on  the  microscope 
slide  and  allowing  the  mixture  to  dry  around  the  edge.     Hemoglobin  crys- 
tals may  be  observed  under  the  microscope  to  have  formed  where  the  solu- 
tion has  concentrated  and  dried. 

Crystals  of  seralbumin  and  of  lactalbumin  can  be  obtained  in  essen- 
tially the  same  manner  already  described  for  ovalbumin. 


106  THE    CHEMICAL    COMPOSITION    OF    THE    BODY 

> 

2.  Elementary  Composition  of   the  Proteins. — a.  Nitrogen. — Make 
an  intimate  mixture  of  some  dry  protein,  preferably  a  casein  preparation, 
with  sodalime  and  place  it  in  a  dry  test-tube.     Warm  gently  over  a  Bunsen 
flame.      Hold  a  piece  of  moistened  red  litmus-paper  over  the  mouth  of 
the  test-tube.      The   ammonia  split  off  from  the   protein  will  color  the 
litmus   blue. 

Warm  together  carefully  in  a  dry  test-tube  a  few  particles  of  dry  protein 
and  a  small  cube  of  metallic  sodium.  (Do  not  place  the  tube  in  the  flame.) 
When  the  fusion  is  complete  and  the  tube  is  cooled  somewhat,  plunge  the 
end  into  a  small  amount  of  water  in  a  casserole.  The  glass  of  the  test-tube 
will  probably  break.  When  the  fused  mass  has  dissolved,  filter,  and  to  the 
filtrate  add  a  few  drops  of  ferric  chloride  and  ferrous  sulphate  solution. 
On  acidifying  with  hydrochloric  acid  a  precipitate  of  Prussian  blue  is  ob- 
tained. Sodium  ferricyanide  is  formed  in  the  reaction  and  in  consequence 
gives  a  Prussian  blue  with  the  excess  of  iron  present. 

b.  Carbon. — Place  some  desiccated  casein  in  the  end  of  a  piece  of  glass 
tubing,  tap  it  down  gently  so  that  the  lumen  of  the  tube  will  not  be  obstructed. 
Heat  the  casein  gently  over  a  small  Bunsen  flame,  inclining  the  tube  so  that 
there  will  be  a  slight  current  of  air  passing  upward  through  it.     The  casein 
will  char,  indicating  the  presence  of  carbon. 

c.  Hydrogen. — Note  the  clear  fluid  that  has  condensed  in  the  upper 
portion  of  the  tubing  from  the  preceding  experiment.     If  a  little  anhydrous 
copper  sulphate  is  introduced  into  the  tube,  the  fluid  of  condensation  coming 
in  contact  with  it  will  become  blue,  indicating  that  water  has  been  formed 
in  the  charring  of  the  casein.     Hydrogen  in  protein  has  been  oxidized  to  form 
water. 

d.  Sulphur. — Boil  some  casein  or  some  egg-white  solution  with  sodium 
hydroxide  after  adding  a  few  drops  of  lead  acetate  solution.     The  presence 
of  sulphur  is  shown  by  the  formation  of  a  black-lead  sulphide.     On  adding 
hydrochloric  acid  the  lead  sulphide  formed  will  be  decomposed  and  the  odor 
of  hydrogen  sulphide  will  be  noted. 

e.  Phosphorus. — Heat  some  casein,  preferably  in  a  nickel  crucible,  with 
a  fusion  mixture  composed  of  three  or  four  parts  of  caustic  soda  and  one  part 
of  potassium  nitrate,  warming  cautiously  until  the  mass  becomes  colorless. 
Dissolve  the  residue  when  cool  in  a  small  volume  of  water,  neutralize  and 
acidify  with  nitric  acid  slightly,  and  add  about  5  c.c.  of  ammonium  molyb- 
date  solution.     Warm  for  some  minutes  at  80°.     A  yellow  precipitate  of 
ammonium  phosphomolybdate  is  obtained. 

3.  Color  Reactions  of  the  Proteins. — a.  Millon's  Reaction. — To  about 
5  c.c.  of  a  dilute  solution  of  egg  albumin  in  a  test-tube  add  a  few  drops  of 
Millon's  reagent.     A  white  precipitate  forms  which  turns  red  when  heated. 
This  test  can  be  used  with  advantage  on  solid  proteins.     In  this  case  the 


COLOR  REACTIONS  OF  THE  PROTEINS  1 07 

reagent  is  added  to  suspensions  of  the  solid  substance.  Such  proteins  as  are 
not  precipitated  by  the  mineral  acids  yield  a  red  solution  instead  of  a  red 
precipitate.  Millon's  reagent  consists  of  one  part  mercury  dissolved  in  two 
parts  by  weight  of  concentrated  nitric  acid;  the  resulting  solution  is  diluted 
with  two  volumes  of  water. 

This  reaction  is  due  to  the  presence  of  the  hydroxyphenyl  group  or 
C6H5OH  in  the  protein  molecule.  Accordingly,  certain  non-protein  sub- 
stances give  this  reaction;  i.e.,  tyrosine,  phenol  (carbolic  acid),  thymol,  etc. 
The  reaction  given  by  the  protein  is  due  to  the  presence  of  the  amino  acid 
tyrosine,  and  it  is  evident  that  the  test  is  really  an  indication  of  the  presence 
of  the  tyrosine  complex  in  the  protein  molecule. 

b.  Xanthoproteic  Reaction. — To  2-3  c.c.  of  egg-albumin  solution  or  of 
some  dry  casein  in  the  test-tube,  add  concentrated  nitric  acid  and  heat  until 
the  protein  dissolves,  forming  a  yellow  solution.     Cool  the  solution  and  care- 
fully add  ammonium  hydroxide  in  excess.     The  yellow  color  changes  to  an 
orange.     This  reaction  is  due  to  the  presence  in  the  protein  molecule  of  the 
phenyl  group  in  phenyalanine,  tyrosine,  or  tryptophane;  with  the  phenyl 
group  nitric  acid  forms  certain  nitro-derivatives  of  benzene. 

c.  Adamkiewicz  or  Hopkins-Cole  Reaction. — Mix  a  couple  of  c.c.  of  con- 
centrated sulphuric  acid  with  4  or  5  c.c.  of  glacial  acetic  acid  in  a  test-tube. 
Add  a  few  drops  of  egg-albumin  solution  and  warm  gently.     A  reddish- 
violet  color  is  produced.     This  reaction  is  due  to  the  presence  of  trypto- 
phane in  the  protein  and  the  test  depends  on  the  presence  of  glyoxylic  acid 
(CHOH2COOH)  in  the  acetic  acid.     The  Hopkins-Cole  reagent,  a  glyoxylic 
acid  solution,  may  be  used  instead  of  the  glacial  acetic  acid.     The  reagent 
is  prepared  by  adding  sodium  amalgam  to  a  saturated  solution  of  oxalic 
acid  and  allowing  the  mixture  to  stand  until  the  evolution  of  gas  ceases. 
In  making  the  test  the  protein  solution  and  Hopkins-Cole  reagent  are  mixed 
thoroughly  in  a  test-tube,  and  concentrated  sulphuric  acid  poured  gently 
into  the  tube  which  has  been  inclined  somewhat  so  that  it  forms  a  layer  in 
the  bottom  of  the  test-tube.     The  acid  and  protein  solutions  will  be  strati- 
fied and  a  reddish-violet  ring  is  developed  where  the  two  fluids  come  in 
contact. 

This  reaction  is  due  to  the  presence  of  tryptophane  in  the  protein  mole- 
cule. Gelatin  does  not  respond  to  this  test,  for  it  does  not  yield  this  sub- 
stance as  a  cleavage  product. 

d.  Liebermann's  Reaction. — Add  a  few  drops  of  egg-white  solution  or  a 
little  dry  casein  to  about  5  c.c.  of  concentrated  hydrochloric  acid  in  a  test- 
tube  and  boil  the  mixture  until  a  pinkish-violet  color  results.     It  was  for- 
merly thought  that  this  reaction  indicated  the  presence  of  a  carbohydrate 
group  in  the  protein  molecule,  but  this  is  now  considered  uncertain. 

e.  Biuret  Reaction. — To  2  or  3  c.c.  of  egg-white  solution  in  a  test-tube  an 


108  THE    CHEMICAL    COMPOSITION    OF    THE   BODY 

equal  volume  of  concentrated  potassium  hydroxide  solution  is  added  and 
mixed  thoroughly.  Very  dilute  (2  per  cent.)  copper  sulphate  solution  is 
added  until  a  purplish-violet  or  pinkish-violet  color  is  produced.  This 
reaction  is  given  by  substances  containing  twoamino  groups  in  the  molecule, 
these  groups  being  joined  directly  together  or  through  a  single  atom  of 
nitrogen  or  carbon.  Non-protein  substances  that  contain  the  necessary 
groups  will  of  course  respond  to  this  test,  which  derives  its  name  from  the 
fact  that  it  is  given  by  biuret,  a  substance  formed  on  heating  urea  to  180°. 

NH2 

I 
NH2  CO 

2C=0         =       NH        +       NHS 
\  \ 

NH2  CO 

NH2 

Urea.  Biuret.       Ammonia 

Proteins  give  this  reaction  since  there  are  more  than  one  CONH2  group 
fn  the  protein  molecule.  Proteoses  and  peptones  give  a  pink  biuret  re- 
action, gelatin  a  rather  blue  reaction,  and  the  ordinary  proteins  a  purple. 

/.  Molisch  Reaction. — This  reaction  is  really  a  carbohydrate  test,  but  is 
given  by  some  proteins  and  interpreted  as  indicating  that  such  proteins 
contain  a  carbohydrate  moiety.  The  test  is  made  as  follows : 

Place  about  5  c.c.  of  the  solution  to  be  tested  in  a  test-tube,  and  add  a 
couple  of  drops  of  a  15  percent,  alcoholic  solution  of  a-naphthol.  Incline 
the  tube  and  pour  very  carefully  down  the  side  about  5  c.c.  of  concentrated 
sulphuric  acid  so  that  the  two  solutions  are  stratified.  A  blue  or  violet-red 
ring  is  obtained  in  the  area  of  contact  of  the  solutions. 

4.  Precipitation  Reactions  of  the  Proteins. — a.  Precipitation  with 
Concentrated  Mineral  Acids. — Prepare  four  test-tubes  which  contain  about 
5  c.c.  of  egg-white  solution.  To  these  respectively  add  drop  by  drop  con- 
centrated sulphuric  acid,  hydrochloric  acid,  nitric  acid,  and  acetic  acid. 
Note  that  the  mineral  acids  precipitate  the  protein.  The  precipitation  with 
nitric  acid  is  a  frequently  used  protein  reaction,  and  when  carried  out  as 
follows  is  known  as  Heller's  ring  test.  The  solution  to  be  tested  is  placed 
in  a  test-tube,  the  tube  is  inclined  and  about  5  c.c.  of  concentrated  nitric 
acid  is  poured  carefully  down  the  side  of  the  tube  so  that  the  solution  and 
acid  stratify.  A  white  zone  of  precipitated  protein  is  obtained  between  the 
strata.  An  instrument  known  as  the  albumiscope  has  been  devised  to 
facilitate  the  making  of  the  ring  tests.  Heller's  ring  test  is  most  commonly 
used  to  determine  the  presence  of  protein  in  urine. 


PRECIPITATION    REACTIONS    OF    THE    PROTEINS  IOQ 

b.  Precipitation  with  Heavy  Metals. — Proteins  form  insoluble  compounds 
with  the  metals  when  mercuric  chloride,  lead  acetate,  copper  sulphate, 
silver  nitrate,  etc.,  are  added  to  protein  solutions. 

c.  Acetic  Acid  and  Potassium  Ferrocyanide  Test. — To  about  5  c.c.  of  egg- 
white  solution  in  a  test-tube  add  five  to  ten  drops  of  acetic  acid  and  then 
potassium  ferrocyanide   drop   by   drop   until   a   precipitate   forms.     This 
test  is  very  delicate. 

d.  Precipitation  with  the  Alkaloidal  Reagents. — Prepare  six  tubes  contain- 
ing about  3  c.c.  egg-white  solution.     To  the  first  add  picric  acid  drop  by 
drop  until  excess  of  the  reagent  has  been  added,  noting  the  changes  with 
care.     Repeat  the  experiment  with   trichloracetic   acid  and  tannic   acid. 
Acidify  the  remaining  tubes  with  hydrochloric  acid,  and  repeat  the  experi- 
ment with   phosphotungstic  acid,  phosphomolybdic   acid,   and   potassium 
mercuric  iodide. 

e.  Heat  Coagulation. — Take  about  10  c.c.  of  egg-white  solution  in  a  test- 
tube  and  heat  to  boiling.     Then  add  a  few  drops  of  dilute  acetic  acid.     The 
protein  will  be  coagulated.     The  acetic  acid  should  be  added  after  heating, 
since    otherwise   acid   metaprotein   might  be   formed.     The   presence    of 
some  neutral  inorganic  salts  tends  to  give  a  sharper  test.     The  addition  of 
the  acid  also  will  dissolve  the  earthy  phosphates  which  are  often  precipi- 
tated from  the  urine  on  heating.     Proteoses,  peptones,  the  casein  of  milk, 
and  a  few  other  proteins  are  not  coagulated  by  heat. 

/.  Precipitation  by  Alcohol. — Add  some  95  per  cent,  alcohol  to  a  test-tube 
containing  about  3  c.c.  of  egg-white  solution.  The  protein  is  precipitated, 
and  on  standing  it  is  coagulated  so  that  it  can  no  longer  be  dissolved  in 
neutral  solvent. 

g.  Salting-out  Experiments. — Add  to  some  diluted  blood-serum  in  a  small 
beaker,  crystals  of  magnesium  sulphate  until  no  more  of  the  salt  will  go 
into  solution.  After  standing  for  a  few  minutes,  filter  and  test  the  filtrate 
and  residue  for  protein  by  some  of  the  precipitation  or  color  reactions  given 
above.  It  is  found  that  the  filtrate  still  contains  some  protein.  Further, 
that  this  protein  can  be  precipitated  on  adding  a  few  drops  of  dilute  acetic 
acid.  When  the  blood  serum  is  similarly  saturated  with  ammonium 
sulphate,  there  will  be  no  protein  found  in  the  filtrate.  If  to  the  blood 
serum  an  equal  volume  of  saturated  ammonium  sulphate  solution  is 
added,  the  result  will  be  the  same  as  that  already  obtained  with  mag- 
nesium sulphate.  Some  proteins  then  are  precipitated  on  saturating 
their  solutions  with  magnesium  sulphate  or  by  adding  an  equal  volume 
of  saturated  ammonium  sulphate  solution;  albumins,  globulins,  and 
proteoses,  however,  are  all  precipitated  by  saturation  with  the  more 
soluble  ammonium  sulphate. 


110 


THE    CHEMICAL    COMPOSITION    OF    THE    BODY 


ALBUMINS  AND  GLOBULINS. 

5.  Properties  of  Albumins  and  Globulins. — Try  out  the  solubility 
and  the  precipitation  tests  indicated  in  the  following  table  for  a  solution  of 
ovalbumin,  and  on  edestin  furnished  by  the  instructor. 


Soluble  in 

Precipitated 

Coagulated 

by 

g 

<n 

fi* 

<n 

Protein 

is 
1 

e 

£ 

1 
1 
? 

Very  dilute  acid 

1 
Q 

On  saturation 
with  NaCl* 

On  saturation 
with  MgSO4* 

On  half-saturatif 
with  (NH  4)2804" 

On  saturation 
with  (NH4)2SO 

Alcohol  in  exces 

1 

Alcohol  J 

Albumin 

(ovalbumin) 

Globulin 

(edestin) 

ALBUMINOIDS. 

6.  Keratin. — Horn  shavings  are  most  conveniently  used  for  the  experi- 
ments with  keratin. 

a.  Try  the  solubility  of  keratin  in  water,  10  per  cent,  salt  solution,  dilute 
hydrochloric  acid,  and  dilute  potassium  or  sodium  hydroxide. 

b.  Make  a  test   for   loosely   combined   sulphur  as  in  experiment  2,  d, 
page  1 06. 

c.  Try  Millon's  reaction  and  the  biuret  reaction,  putting  the  undissolved 
shavings  directly  into  the  reagent. 

d.  Try  the  solubility  of  keratin  in  the  artificial  gastric  and  pancreatic 
juice  furnished  by  the  instructor. 

7.  Collagen. — The  tendo  achillis  of  the  ox  may  be  used  for  the  prepara- 
tion of  collagen. 

a.  Clean  the  tendon  and  cut  it  into  small  pieces.     Wash  the  pieces  in 
dilute  salt  solution  in  order  to  remove  the  soluble  protein,  and  then  wash 
with  distilled  water.     Transfer  the  pieces  of  washed  tendon  to  a  flask  and 
add  100  c.c.  of  saturated  lime-water  and  same  amount  of  distilled  water. 
The  flask  should  be  shaken  at  intervals  for  twenty-four  hours.     The  lime- 
water  dissolves  the  mucoid  in  the  tendon.     Filter  off  the  pieces  of  tendon 
and  save  the  filtrate  for  a  later  experiment.     The  residue  of  the  tendon  con- 
sists of  the  albuminoid  collagen  and  a  little  elastin.     We  may  consider  the 
tests  to  follow  as  being  made  upon  collagen. 

b.  Cut  the  collagen  into  very  fine  pieces  and  try  the  solubility  as  in  6  a 
above. 

*  If  not  precipitated,  acidify  and  note  result. 

t  See  page  89. 

J  On  standing  under  alcohol. 


GLYCOPROTEINS  III 

c.  Test  for  loosely  combined  sulphur  in  collagen.     Heat  with  the  alkali 
until  the  large  piece  of  collagen  used  is  partly  decomposed,  then  add  one  or 
two  drops  of  lead  acetate  and  again  heat  to  boiling. 

d.  Try  Millon's  reaction,  the  biuret  test,  the  xanthoproteic  reaction,  and 
the  Hopkins-Cole  reaction. 

e.  Formation  of  gelatin  from  collagen.     Transfer  the  remainder  of  the 
collagen  to  a  casserole,  fill  about  two-thirds  with  water,  and  boil  for  several 
hours,  replacing  enough  of  the  water  that  is  lost  by  evaporation  to  keep  the 
pieces  well  covered.     Filter  and  cool.     Collagen  is  transformed  into  gelatin 
and  the  characteristic  gel  is  obtained.     Try  this  experiment  also  with  some 
cartilage  from  the  trachea  and  with  some  0.2  per  cent.  HC1  extracted  bone. 

/.  The  properties  of  gelatin.  Try  the  solubility  as  in  a  above  on  some 
gelatin  furnished  by  the  instructor.  Try  the  solubility  also  in  hot  water. 

g.  Try  the  tests  for  loosely  combined  sulphur.  Try  Millon's  Hopkins- 
Cole,  and  the  biuret  reactions.  Try  the  solubility  in  artificial  gastric  and 
pancreatic  juices. 

8.  Elastin. — a.  Preparation  of  Elastin. — Cut  the  ligamentum  nuchce  of, 
the  ox  into  strips  and  run  it  through  a  meat  chopper.     Wash  the  finely 
divided  material  with  salt  solution  and  in  running  water  for  some  hours. 
Transfer  it  to  a  flask  and  add  about  200  c.c.  of  half-saturated  lime-water  and 
extract  for  forty-eight  hours.     Filter  off  the  lime-water  which  has  dissolved 
the  mucoid  present,  saving  the  filtrate,  and  wash  the  residue  and  tendon  with 
water.     Boil  the  material  in  dilute  acetic  acid  for  some  hours  to  convert  the 
collagen  present  into  gelatin.     Wash  the  residue  thoroughly  with  water.     It 
may  be  dried  by  washing  it  with  boiling  alcohol,  pressing  it  out  between 
filters,  and  manipulating  it  in  the  air. 

b.  Try  the   solubility   of  elastin   as   in  a.     Test  for  loosely  combined 
sulphur.     Try  Millon's,  biuret,  and  the  Hopkins-Cole  reactions.     Try  the 
solubility  in  artificial  gastric  and  pancreatic  juices. 

c.  Make  a  table  comparing  the  properties  of  keratin,  collagen,  and  elastin. 

GLYCOPROTEINS. 

9.  Mucoid. — a.  Unite  the  lime-water  of  filtrates  obtained  in  the  extraction 
of  the  tendons  in  the  preceding  experiment  and  acidify  them  with  dilute 
acetic  acid  until  precipitation  occurs.     The  mucoid  is  precipitated.     Avoid 
adding  excessive  acid  or  the  mucoid  will  again  go  into  solution.     Allow  the 
mucoid  precipitate  to  settle  and  decant  the  supernatant  fluid  and  filter  off 
the  precipitate. 

b.  Try  the  biuret  test  on  a  portion  of  the  mucoid. 

c.  Place  the  remainder  of  the  mucoid  in  a  beaker,  add  about  25  c.c.  of 
water  and  2  c.c.  of  dilute  hydrochloric  acid;  boil  until  the  solution  becomes 
dark.     To  a  portion  of  the  mixture  add  a  few  drops  of  barium  chloride.     A 


112  THE    CHEMICAL    COMPOSITION    OF    THE    BODY 

white  precipitate  shows  the  presence  of  sulphate.  Neutralize  to  litmus-paper 
with  solid  potassium  or  sodium  hydroxide  and  test  for  sugar  with  Fehling's 
solution  (see  Experiment  12,  d).  The  mucoid  has  been  split  by  boiling  with 
acid  into  protein,  protein  cleavage  products,  and  carbohydrate  or  a  reducing 
substance.  The  presence  of  oxidized  sulphur  in  the  molecule  has  also 
been  shown. 

'DERIVED  PROTEINS. 

10.  Metaproteins. — a.   Acid   Metaproteins  or  Acid   Albuminate. — Acid 
metaprotein  or  acid  albumin  may  be  prepared  as  follows: 

Dilute  solution  of  white  of  egg  with  several  volumes  of  .4  per  cent,  hydro- 
chloric acid.  Allow  it  to  stand  for  some  time  at  40°  Centigrade.  Filter  and 
neutralize  the  filtrate.  A  precipitate  of  acid  metaprotein  is  obtained.  The 
acid  albuminate  then  is  insoluble  in  neutral  dilute  salt  solutions,  but  it  dis- 
solves in  acidified  solutions.  Filter  off  some  of  the  precipitated  acid  albumin- 
ate  and  test  it  for  loosely  combined  sulphur. 

b.  Alkali  metaprotein  is  formed  on  treating  proteins  with  alkali.  To 
some  undiluted  egg  white  in  an  evaporating  dish  add  sodium  hydroxide  solu- 
tion slowly  with  constant  stirring.  The  mixture  forms  a  stiff  gel  known  as 
Lieberkuhn's  jelly.  Wash  the  gel,  which  has  been  broken  into  small  pieces, 
with  running  water  until  the  excess  of  alkali  is  removed.  Warm  it  in  a  small 
amount  of  water  and  dissolve  by  heating  gently.  Neutralize  the  solution 
carefully  with  acid,  noting  the  odor  of  the  hydrogen  sulphide  that  is  given  off. 
The  precipitate  which  appears  when  the  neutral  point  is  reached  is  alkali 
metaprotein  or  alkali  albuminate.  Filter  off  the  precipitate,  wash  it  in 
water,  and  try  the  test  for  loosely  combined  sulphur.  A  weak  reaction  or 
negative  result  shows  that  the  loosely  combined  sulphur  has  been  split  off  by 
treating  the  protein  with  the  alkali,  a  change  which  has  not  occurred  in  the 
formation  of  the  acid  albuminate  above. 

SECONDARY  PROTEIN  DERIVATIVES. 

11.  Proteose   and    Peptone. — Commercial  proteose- peptone  prepara- 
tions, such  as  Witte  peptone  or  Armour's  peptone,  may  be  employed  for 
the  separation  of  proteoses  and  peptones. 

a.  Take  about  5  grams  of  the  proteose-peptone  mixture  and  dissolve  it  in 
100  c.c.  of  water. 

Try  the  biuret  reaction,  Millon's  reaction,  and  Heller's  ring  test.  Do 
the  proteoses  and  peptones  coagulate  on  heating  ? 

b.  Place  the  remainder  of  the  solution  in  the  beaker  and  add  dry  ammonium 
sulphate  in  excess.     Note  that  before  the  solution  is  completely  saturated  with 
the  salt,  the  precipitation  of  the  primary  proteoses  (proto  proteose  and  hetero 
proteose).     Completely  saturate  the  solution  with  the  ammonium  sulphate, 
warming  it  gently  to  facilitate  the  separation.     At  full  saturation  the  second- 


CARBOHYDRATES  113 

ary  proteoses  (deutero-proteoses)  are  precipitated.  The  peptones  remain 
in  solution.  Try  the  biuret  test  on  the  precipitate  and  on  the  filtrate  con- 
taining the  peptone,  in  the  latter  instance  making  the  solution  strongly  alka- 
line with  solid  sodium  or  potassium  hydroxide. 

d.  From  what  you  have  learned  of  the  properties  of  the  derived  proteins 
in  the  text  and  in  the  laboratory,  prepare  a  chart  (Exp.  5)  in  which  the 
properties  of  these  substances  are  indicated.  Compare  the  results  in  this 
chart  and  the  results  in  this  table  with  the  similar  one  that  you  made  for 
the  albumins  and  globulins 

CARBOHYDRATES. 

12.  Starch. — a.  Examine  under  the  microscope  and  sketch  the  starch 
granules  obtained  by  grinding  some  potato  scrapings  in  a  mortar  with  a 
little  water;  examine  and  sketch  also  the  granules  of  corn,  wheat  (flour), 
and  arrowroot  starch. 

b.  Starch  Paste. — Make  a  suspension  of  i  gram  of  arrowroot  starch  in  a 
little  distilled  water  by  grinding  in  a  mortar,  and  pour  slowly  into  some 
boiling  water.      Heat   for  a  few  minutes   longer,    cool    and  make  up  to 
100  c.c. 

c.  Iodine  Test. — Shake  up  three  or  four  drops  of  dilute  iodine  solution 
with  2  c.c.  starch.     A  deep  blue  color  appears.     The  color  is  discharged  in 
dilute   alkali   and  reappears  on  acidifying   again.     Heat   also  discharges 
the  color. 

d.  Fehling's   Test.— Fill  a  test-tube   about  a  fourth  full  of  Fehling's 
solution  and  heat  to  boiling  for  a  minute  or  two.     Add  a  few  drops  of  starch 
paste.     The  red  precipitate  of  copper  (cuprous)  oxide  should  not  be  obtained, 
for  pure  starch  does  not  reduce  Fehling's  solution. 

e.  Hydrolysis  of  Starch. — Boil  starch  solution  with  5  per  cent,  sulphuric 
acid  for  fifteen  minutes.     Test  with  Fehling's  solution,  first  neutralising  the 
excess  of  acid.     A  copious  precipitate  of  cuprous  oxide  shows  that  the  starch 
has  been  converted  to  reducing  sugar. 

13.  Dextrins. — Make  a  5  per  cent,  solution  of  dextrin  in  distilled  water 
and  try: 

a.  Iodine  Test. — This  gives  a  red  which  is  characteristic. 

b.  Fehling's  Test. 

14.  Dextrose. — Test  a  5  per  cent,  solution  of  dextrose: 

a.  Iodine  Test. — No  reaction. 

b.  Fehling's  Test. 

15.  Glycogen. — Use  i  per  cent,  solution  of  glycogen.     Note  the  char- 
acteristic opalescence  of  the  solution. 

a.  Iodine  Test. — A  wine-red,  somewhat  like  that  given  by  dextrin. 

b.  Fehling's  Test. — Glycogen  does  not  reduce  the  copper  solution. 

c.  Hydrolysis. — Test  as  in  e.  The  glycogen  is  hydrolyzed  to  dextrose. 
8 


114  THE   CHEMICAL   COMPOSITION   OF   THE    BODY 

FATS. 

16.  Neutral  Fat. — a.  Melting-point. — Compare  neutral  olive  oil,  some 
fresh  rendered  lard,  and  some  tallow.     The  former  is  fluid  at  ordinary 
room  temperature.     Determine  the  melting-points  of  the  lard  and  of  the 
tallow  by  the  method  of  Wiley.     Fill  a  test-tube  one-half  full  of  water  and 
add  a  two-inch  top  layer  of  alcohol.     Prepare  a  thin  flake  of  fat  and  suspend 
it  in  the  test-tube  at  the  dividing  line  of  the  water  and  alcohol.     Insert  the 
bulb  of  a  thermometer  at  the  same  level.     Mount  the  test-tube  with  the 
thermometer  in  a  beaker  on  a  ring  stand,  fill  the  beaker  with  water  above  the 
level  of  the  content  of  the  test-tube,  and  gradually  heat  with  stirring  of  the 
water  in  the  beaker.     At  the  melting  temperature  the  flake  of  fat  will  run 
into  a  round  drop. 

b.  Solubility. — Fat  is  insoluble  in  water,  but  soluble  in  acetone,  ether, 
chloroform,  benzol,  and  in  alcohol. 

c.  Saponification. — Heat  some  fat  in  an  evaporating-dish,  add  sodium 
hydrate,  and  boil.     Saponification  takes  place.     The  soap  is  soluble  in  water. 
Add  25  per  cent,  sulphuric  acid  to  some  of  the  soap,  the  fatty  acid  is  liberated 
and  collects  on  the  surface  of  the  solution. 

17.  Fatty  Acids. — Collect  some  of  the  fatty  acids,  wash  to  remove 
excess  of  sulphuric  acid,  and  dissolve  in  ether. 

a.  Acid  Reaction. — Add  an  ether  solution  of  the  fatty  acid  to  neutral 
litmus,  or  to  faintly  alkaline  phenolphthalein.     The  former  turns  red,  and 
the  red  of  the  alkaline  solution  of  the  latter  is  discharged. 

b.  Acrolein  Test. — Evaporate  the  ether  from  2  c.c.  of  the  solution,  add 
potassium  bisulphate  crystals  to  the  acid  in  a  test-tube,  and  raise  to  a  high 
heat  over  a  Bunsen.     No  acrolein  is  given  off.     Repeat  on  neutral  fat  and 
on  glycerin.     Both  liberate  the  irritating  fumes  of  acrolein. 

18.  Emulsification. — a.  Shake    up    neutral    olive    oil    and    water,    no 
emulsion  is  formed  and  the  oil  quickly  separates. 

b.  Add  a  couple  of  drops  of  fatty  acid,  a  very  good  but  temporary  emul- 
sion is  now  formed. 

c.  Use  rancid  fat,  a  temporary  emulsion  is  formed. 

d.  Add  a  little  soap  to  each  of  the  above.    A  good  permanent  emulsion 
is  now  formed,  but  best  in  c. 

19.  Lipoids. — a.  Grind    some    pig's    or   calf's  brain  with  ether  in  a 
mortar,  place  in  a  flask  with  sufficient  ether  to  make  a  thin  suspension  of 
the  material  and  set  aside  until  the  undissolved  material  has  sedimented 
completely  to  the  bottom.     Decant  the  clear  ether  and  add  acetone  until 
the  lecithins  have  been  precipitated.     Cholesterol  will  crystallize  from  the 
filtrate   on  spontaneous  evaporation  of   the  latter.     Sketch  the  cholesterol 
crystals. 

b.  Show  the  presence  of  nitrogen  in  lecithin  (see  Exp.  2  a). 


FATS  115 

;.  Try  the  acrolein  test  as  in  Exp.  17  b. 

d.  Fuse  some  lecithin  in  a  metal  crucible  with  a  fusion  mixture  of  3 
parts  of  caustic  potash  and   i   part  of  potassium  nitrate.     Dissolve  in  a 
small  volume  of  water,  acidify  slightly  with  nitric  acid,  add  a  few  drops  of 
ammonium  molybdate  solution,  and  warm  to  75°  C.  for  a  few  minutes.     A 
yellow  precipitate  of  ammonium  phosphomolybdate  indicates  the  presence 
of  phosphorus  in  the  lecithin. 

e.  Try  to  saponify  some  cholesterol  as  in  Exp.  16  c.     As  cholesterol  is 
not  a  fat,  saponification  does  not  take  place. 

20.  The  Salts. — Coagulate  the  protein  in  10  grams  of  chopped  meat  or 
25  c.c.  of  blood  by  boiling  for  a  few  minutes  with  25  c.c.  of  water  to  which  a 
few  drops  of  acetic  acid  have  been  added.  Filter  off  the  coagulated  protein, 
wash  the  precipitate  on  the  paper  with  a  very  little  hot  water,  and  add  the 
washings  to  the  original  filtrate.  The  coagulum  should  filter  off  quickly 
and  the  filtrate  should  be  perfectly  clear;  otherwise  repeat  the  experiment. 
Make  the  following  tests: 

a.  Chlorides. — Acidify  a  small  portion  of  the  filtrate  with  nitric  acid 
and  add  a  few  drops  of  silver  nitrate  solution.     A  white  precipitate,  which 
dissolves  on  adding  ammonia  and  reappears  on  acidification  with  nitric 
acid,  shows  the  presence  of  chlorides. 

b.  Sulphates. — Acidify  a  portion  of  the  filtrate  with  hydrochloric  acid 
and  add  a  few  drops  of  barium  chloride  solution.     A  white  precipitate  of 
barium  sulphate  indicates  that  sulphates  are  present.     A  much  stronger 
reaction  can  be  obtained  when  the  ash  or  alkali  fusion  of  a  tissue  is  tested, 
the  sulphur  in  the  protein  then  having  been  oxidized  to  a  sulphate. 

c.  Phosphates. — Acidify  with  nitric  acid  and  then  add  a  few  drops  of 
ammonium  molybdate  solution.     Warm  in  the  water-bath  at  75°  C.  for  a 
few  minutes.     A  yellow  precipitate  of  ammonium  phosphomolybdate  indi- 
cates the  presence  of  phosphate. 

d.  Calcium. — Add  a  few  drops  of  ammonium  oxalate  solution  to  a  por- 
tion of  the  filtrate.     A  white  precipitate  of  calcium  oxalate  forms.     On 
microscopic  examination  this  is  found  to  consist  of  characteristic  minute 
octahedrals. 

e.  Iron. — Acidify  with  hydrochloric  acid  and  add  a  few  drops  of  potas- 
sium ferrocyanide.     A  Prussian  blue  color  indicated  the  presence  of  iron. 
Try  the  reaction  on  some  blood  which  has  been  ashed  in  a  crucible  and 
dissolved  out  in  a  little  dilute  hydrochloric  acid.     The  iron  is  present  in  the 
compound  protein,  hemoglobin,  of  the  red  corpuscles. 

/.  Magnesium. — Tease  out  some  fibers  of  frog  muscle  in  a  few  drops  of 
water  on  a  microscope  slide  and  invert  a  beaker  containing  a  piece  of  filter- 
paper  moistened  with  ammonia  over  it.  After  a  few  minutes,  examine 
under  the  microscope.  Characteristic  star-shaped  or  fern-like  crystals  of 
ammonium  magnesium  phosphate  will  be  seen. 


PLATE  II 

VARIETIES  OF  LEUCOCYTES 

Plate  II  is  reproduced  by  the  kind  permission  of  Dr.  Cabot.  It  illustrates  certain 
typical  varieties  of  leucocytes.  All  are  stained  with  the  Ehrlich  triacid  stain,  and  drawn 
with  camera  lucida.  Oil  immersion  objective  ^  and  ocular  No.  iii  of  Leitz.  (Cabot.) 

1.  Small  Lymphocytes. — In  the  cell  at  the  left  note  the  transparent  protoplasm; 
in  the  cell  next  to  it  note  the  very  pale  pink  of  protoplasm  around  the  nucleus  which 
is  deeply  stained,  especially  at  the  periphery.     The  next  cell  has  an  indented  nucleus; 
its  protoplasm  relatively  distinct.     The  cell  on  the  extreme  right  shows  no  protoplasm 
and  is  probably  necrotic.     In  all  note  absence  of  granules  with  this  stain.     With  basic 
stains  a  blue  network  appears  in  the  protoplasm. 

2.  Large  Lymphocytes. — Note  the  pale  stain  of  nuclei  and  protoplasm,  regularity 
of  outline;  indented  nucleus  in  one.     Every  intermediate  stage  between  these  and  the 
"small"  lymphocytes  occurs,  and  the  distinction  between  them  is  arbitrary. 

3.  Polymorphonuclear  Neutrophiles. — Note  the  varieties  in  size  and  shape  of  granules, 
the  regular  staining  of  the  nuclei,  the  light  space  around  them,  their  relatively  central 
position  in  the  cell. 

.  4.  Myelocytes. — Note  the  identity  of  granules  with  those  just  described;  the  even, 
pale  stain  of  nuclei,  their  position  near  the  surface  (edge)  of  the  cell.  The  two  cells 
figured  indicate  the  usual  variations  in  size  of  the  whole  cell. 

5.  Eosinophile. — Note    regular   shape,    loose  connection  of  granules,  their  copper 
color,  their  uniform  and  relatively  large  size,  and  spherical  shape. 

6.  Eosinophilic  Myelocyte. — Note   similarity  to  the  ordinary  myelocytes  b,  except 
as  regards  granules.     Colors  of  granules  may  be,  as  in  e,  ordinary  eosinophile. 


116 


KIRKES'  PHYSIOLOGY 


PLATE  II 


Polymorphonuclear 

neutrophiles     t_  __^H 


Myelocytes 


— dBHI      Small  Lymphocytes 


——•Large  Lymphocytes 


Eosinophile 


^^HL       Eosinophilic 
Y"*  Myelocyte 


Varieties  of  Leucocytes 


R.  C.  Cabot  fee. 


CHAPTER  IV 

THE  BLOOD 

THE  blood  is  the  fluid  medium  from  which  all  the  tissues  of  the  body  are 
nourished.  By  means  of  the  blood,  materials  absorbed  from  the  alimentary 
canal  as  well  as  oxygen  taken  from  the  air  in  the  lungs  are  carried  to  the 
tissues,  while  substances  which  result  from  the  metabolism  of  the  tissues  are 
carried  to  the  excretory  organs  to  be  removed  from  the  body.  The  blood 
acts  as  a  medium  of  exchange  between  the  various  tissues  themselves.  A 
good  example  is  the  activity  of  the  blood  in  regulating  the  reaction  of 
the  body  in  balanced  neutrality.  The  blood  is  also  an  important  factor 
in  the  regulation  of  the  body  temperature. 

The  blood  is  a  somewhat  viscid  fluid,  and  in  man  and  in  all  other 
vertebrate  animals,  with  the  exception  of  the  two  lowest,  is  red  in  color. 
The  exact  color  of  blood  is  variable;  that  taken  from  the  systemic  arteries, 
from  the  left  side  of  the  heart  and  from  the  pulmonary  veins  is  of  a  bright 
scarlet  hue;  that  obtained  from  the  systemic  veins,  from  the  right  side  of 
the  heart,  and  from  the  pulmonary  artery  is  of  a  much  darker  color,  which 
varies  from  bluish-red  to  reddish-black.  At  first  sight  the  red  color  appears 
to  belong  to  the  whole  mass  of  blood,  but  on  further  examination  this  is 
found  not  to  be  the  case.  In  reality  blood  consists  of  an  almost  colorless 
fluid,  called  plasma  or  liquor  sanguinis,  in  which  are  suspended  numerous 
minute  masses  of  protoplasm,  called  blood  corpuscles.  The  corpuscles  are 
of  the  two  varieties,  the  white  ameboid  corpuscles,  or  leucocytes,  and  the 
red  corpuscles,  erythrocytes.  The  latter  compose  by  far  the  larger  mass  of 
blood-cells.  They  contain  the  red  pigment,  hemoglobin,  to  which  the  color 
of  the  blood  is  due. 

The  plasma  or  fluid  part  of  the  blood  is  a  remarkably  complex  chemical 
mixture.  It  is  kept  in  constant  rapid  circulation  through  the  blood  vessels 
of  the  body  and  is,  therefore,  thoroughly  mixed  and  homogeneous  in 
character. 

Quantity  of  the  Blood. — The  quantity  of  blood  in  any  animal  under 
normal  conditions  bears  a  fairly  constant  relation  to  the  body  weight. 
The  amount  of  blood  in  man  averages  -^  to  -—  of  the  body  weight.  In 
other  mammals  the  proportion  of  blood  is  also  fairly  constant,  varying 
from  ~  to  -^5-  of  the  body  weight.  In  many  of  the  lower  vertebrates,  the 
fishes  for  example,  the  relative  quantity  of  blood  is  very  much  less. 

It  is  difficult  to  make  an  exact  estimate  of  the  quantity  of  blood  in 
animals  or  in  man  though  the  blood  volume  in  man  is  of  great  importance 
from  the  standpoint  of  disease.  Measurements  are  constantly  given 

117 


Il8  THE    BLOOD 

for  the  number  and  distribution  of  the  corpuscles  and  of  the  blood  volume 
index.  This  data,  to  be  of  value,  must  be  compared  with  the  total  pro- 
portionate amount  of  blood.  A  comparatively  recent  determination  of 
this  question  was  made  by  methods  that  are  modern  by  Keith,  Rowntree, 
and  Geraghty.  They  injected  a  known  quantity  of  a  dye  that  is  absorbed 
with  difficulty,  vital  red.  As  soon  as  the  dye  was  distributed,  three 
minutes,  they  withdrew  blood  into  powdered  oxylate  from  the  correspond- 
ing vein  of  the  other  arm,  centrifuged  quickly  and  compared  the  stained 
plasma  obtained  from  the  arm  with  a  standard  dilution  of  the  dye,  using 
the  colorimetric  method.  Computation  yielded  the  following  facts. 
The  total  blood  averaged  8.8  per  cent.,  ~^  of  the  body  weight.  This  is 
about  85  cubic  centimeters  per  kilo.  The  plasma  averaged  about  50 
cubic  centimeters  per  kilo. 

This  quantity  of  blood  is  distributed  in  the  different  parts  of  the  body, 
chiefly  in  the  muscles,  the  liver,  the  heart,  and  larger  blood  vessels,  as 
shown  by  the  following  figures  determined  on  the  rabbit  by  Ranke  (from 
Vierordt) : 

Per  cent. 

Spleen o .  23 

Brain  and  cord 1.24 

Kidney 1.63 

Skin 2.10 

Abdominal  viscera 6.30 

Cartilage 8 .  24 

Heart,  lungs,  and  large  blood  vessels 22.76 

Resting  muscle 29  . 20 

Liver 29.30 

The  normal  blood  volume  varies  somewhat  with  relation  to  the  intake 
of  food  and  drink.  Relative  body  dessication  would  appear  in  extreme 
thirst  or  under  conditions  of  unusual  loss  of  water  without  increasing  the 
intake,  as  in  extreme  perspiration,  diarrheas,  etc. 

COAGULATION  OF  THE  BLOOD. 

The  most  characteristic  property  which  the  blood  possesses  is  that  of 
clotting  or  coagulating.  This  phenomenon  may  be  observed  under  the  most 
favorable  conditions  in  blood  which  has  been  drawn  into  an  open  vessel.  In 
about  two  or  three  minutes,  at  the  ordinary  temperature  of  the  air,  the  surface 
of  the  fluid  is  seen  to  become  semisolid  or  jelly-like,  and  this  change  takes 
place,  in  a  minute  or  two  afterward,  at  the  sides  of  the  vessel  in  which  it  is 
contained  and  then  quickly  extends  throughout  the  entire  mass.  The  time 
which  is  occupied  in  these  changes  is  about  eight  or  nine  minutes.  The 
solid  mass  is  of  exactly  the  same  volume  as  the  previously  liquid  blood,  and 
adheres  so  closely  to  the  sides  of  the  containing  vessel  that  if  the  latter  be 
inverted  none  of  its  contents  escape.  The  solid  mass  is  the  crassamentum 
or  clot.  If  the  clot  be  watched  for  a  few  minutes,  drops  of  a  light,  straw- 
colored  fluid,  the  serum,  may  be  seen  to  make  its  appearance  on  the  surface, 


COAGULATION    OF    THE    BLOOD  1 19 

and,  as  it  becomes  greater  and  greater  in  amount,  to  form  a  complete  super- 
ficial stratum  above  the  solid  clot.  At  the  same  time  the  serum  begins  to 
transude  at  the  ddes  and  at  the  under  surface  of  the  clot,  which  in  the  course 
of  an  hour  or  two  floats  in  the  liquid.  The  appearance  of  the  serum  is  due 
to  the  fact  that  the  clot  contracts,  thus  squeezing  the  fluid  out  of  its  mass. 
The  first  drops  of  serum  appear  on  the  surface  about  eleven  or  twelve  min- 
utes after  the  blood  has  been  drawn;  and  the  fluid  continues  to  transude  for 
from  thirty-six  to  forty-eight  hours. 

The  clotting  of  blood  is  due  to  the  development  in  the  plasma  of  an  in- 
soluble substance  called  fibrin.  This  fibrin  forms  threads  or  strands  through 
the  mass  in  every  direction.  The  strands  adhere  to  each  other  wherever 
they  come  in  contact,  thus  forming  a  very  dense  tangle  and  meshwork  which 
incloses  within  itself  the  blood-corpuscles.  The  clot  when  first  formed, 
therefore,  includes  the  whole  of  the  blood  in  an  apparently  solid  mass,  but 
soon  the  fibrinous  meshwork  begins  to  contract  and  the  serum  is  squeezed 
out.  When  a  large  part  of  the  serum  has  been  squeezed  out  the  clot  is  found 
to  be  smaller,  but  firmer  and  harder,  as  it  is  now  made  up  more  largely  of 
fibrin  and  blood  corpuscles.  Thus  in  coagulation  there  is  a  rearrangement 
of  the  constituents  of  the  blood;  liquid  blood  consisting  of  plasma  and 
blood  corpuscles,  and  clotted  blood  of  serum  and  clot.  These  relations  are 
roughly  shown  in  the  following  diagram: 

Liquid  blood. 


Plasma.  Corpuscles. 


Serum.  Fibrin. 


Clot. 


Clotted  blood. 

The  rapidity  with  which  coagulation  takes  place  varies  greatly  in  different 
animals  and  at  different  times  in  the  same  animal.  Where  coagulation  is 
very  slow  the  red  corpuscles,  which  are  somewhat  heavier  than  plasma, 
often  have  time  to  settle  considerably  before  the  fibrin  is  formed.  If  the 
blood  is  rapidly  cooled  to  a  temperature  approaching  o°  C.  then  the  clot  is 
very  greatly  delayed.  Horse's  blood  is  particularly  favorable  for  demon- 
strating this  point.  In  it  clotting  occurs  so  slowly  that  very  often  the  red 
corpuscles  will  completely  settle  out,  and  when  the*  blood  is  again  warmed 
and  the  clotting  takes  place  there  is  a  superficial  stratum  differing  in  appear- 
ance from  the  rest  of  the  clot,  having  a  grayish-yellow  color.  This  is  known 


120 


THE   BLOOD 


as  the  bufy  coat  or  crusta  phlogistica.  The  buffy  coat,  produced  in  the  man- 
ner just  described,  commonly  contracts  more  than  the  rest  of  the  clot,  on 
account  of  the  absence  of  colored  corpuscles  from  its  meshes.  When  the 
clot  is  allowed  to  stand,  the  white  corpuscles  which  have  escaped  the  clot 
by  ameboid  movement  settle  on  its  surface  often  in  such  numbers  that  they 
form  a  distinct  superficial  layer,  grayish-white  in  appearance. 

That  the  clotting  of  blood  is  due  to  the  gradual  appearance  in  it  of  fibrin 
may  be  easily  demonstrated,  For  example,  if  recently  drawn  blood  be 
whipped  with  a  bundle  of  twigs  or  wires,  the  fibrin  may  be  withdrawn  from 
the  blood  before  it  can  entangle  the  blood  corpuscles  within  its  meshes. 
It  adheres  to  the  twigs  in  stringy  threads  relatively  free  from  corpuscles. 
The  blood  from  which  the  fibrin  has  been  withdrawn  no  longer  exhibits  the 
power  of  spontaneous  coagulability  and  it  is  now  called  defibrinated  blood. 
Although  these  facts  have  long  been  known,  the  closely  associated  problem 
as  to  the  exact  manner  in  which  fibrin  is  formed  is  by  no  means  so  simple. 


FlG.  107. — Reticulum  of  Fibrin,  from  a  Drop  of  Human  Blood,  after  Treatment  with 

Rosanilin.     (Ranvier.) 

Fibrin  is  derived  from  the  plasma.  Pure  plasma  may  be  procured  by 
delaying  coagulation  in  blood  by  keeping  it  at  a  temperature  slightly  above 
freezing-point,  until  the  colored  corpuscles  have  subsided  to  the  bottom  of 
the  containing  vessel.  The  blood  of  the  horse  is  specially  suited  for  the 
purposes  of  this  experiment.  A  portion  of  the  colorless  supernatant  plasma, 
if  decanted  into  another  vessel  and  exposed  to  the  ordinary  temperature  of 
the  air,  will  coagulate,  producing  a  clot  similar  in  all  respects  to  blood  clot, 
except  that  it  is  colorless  from  the  absence  of  red  corpuscles.  If  some  of 
the  plasma  be  diluted  with  twice  or  three  times  its  bulk  of  normal  saline 
solution  (0.9  per  cent.),  coagulation  is  delayed,  and  the  stages  of  the  gradual 
formation  of  fibrin  in  it  may  be  conveniently  watched.  The  viscidity  which 
precedes  the  complete  coagulation  may  be  actually  seen  to  be  due  to  the 
formation  of  fibrils  of  fibrin — first  of  all  at  the  edge  of  the  fluid-containing 


THEORIES    OF    COAGULATION 


121 


vessel,  and  then  gradually  extending  throughout  the  mass.  If  a  portion  of 
plasma,  diluted  or  not,  be  whipped  with  a  bundle  of  twigs  or  wire  during  the 
process  of  clotting,  the  fibrin  will  be  obtained  as  a  stringy,  insoluble  mass,  just 
in  the  same  way  as  from  the  entire  blood.  The  resulting  fluid  no  longer 
retains  its  powei  of  spontaneous  coagulability  and  is  in  fact  now  a  typical 
serum. 

Theories  of  Coagulation. — It  is  evident  that  the  blood  plasma  contains 
some  substance  or  substances  which  take  part  in  the  formation  of  fibrin. 
By  numerous  investigations  it  has  been  found  that  the  direct  antecedent  of 
the  fibrin  is  the  protein  substance,  fibrinogen.  This  fibrinogen  exists  in  the 
blood  plasma  at  all  times,  but  is  somewhat  increased  under  certain  condi- 
tions. The  fibrinogen  is  reacted  on  by  another  substance  known  as  thrombin. 
We  shall  not  present  the  numerous  theories  which  have  been  held  concerning 
blood  coagulation,  many  of  which  have  been  more  or  less  disproven,  but  shall 
try  to  present  the  condensed  statement  of  the  present  explanations  of  this 

Blood  Tissue  Cell 


Neutral  Salts  Fibrinogen  Calcium  Salts 

(for  dissolving 
fibrinogen) 


Fibrin-globulin 


Thrombok  ina  se 


Fibrin 
Morawitz'  Schema  of  Coagulation. 

intricate  phenomenon.  One  may  start  from  the  statement  that  the  fibrinogen 
of  the  plasma  when  acted  upon  by  the  thrombin,  also  of  the  plasma,  produces  an 
insoluble  substance,  fibrin.  The  chief  interest  centers  around  the  origin  and 
character  of  ihefirbinogen,  the  origin  and  nature  of  the  thrombin,  and  the  con- 
ditions which  influence  its  activity. 

The  fibrinogen  is  present  in  blood  plasma  of  the  circulating  blood  of  the 
body  at  all  times.  It  can  be  separated  from  plasma  by  various  chemical 
means, and  when  purified  can  be  made  to  form  fibrin  under  proper  conditions. 
All  observers  are  agreed  that  this  protein  is  the  immediate  precursor  of  the 
insoluble  fibrin.  Its  origin  in  the  blood  has  been  traced  by  Matthews  to  the 
disintegration  of  the  white  blood  corpuscles. 

The  thrombin  is  the  substance  which  reacts  on  the  fibrinogen  in  the 


122  THE   BLOOD 

processes  of  fibrin  formation.  It  does  not  exist  in  the  living  blood 
vessels,  or  at  least  is  present  only  in  minute  quantities,  but  makes  its 
appearance  immediately  the  blood  is  drawn. 

The  sources  of  these  substances  and  the  part  taken  by  each  in  the 
process  of  coagulation  are  given  by  Morawitz.  If  blood  be  drawn,  centri- 
fugalized,  and  the  leucocytes  and  blood  plates  separated  from  the  plasma 
and  suspended  in  water,  their  solution  will  cause  the  formation  of  fibrin 
from  fibrinogen  in  the  presence  of  calcium.  The  leucocytes  and  platelets 
are,  therefore,  the  source  of  thrombin.  The  thrombin  can  be  isolated  in  a 
stable  condition,  and  when  its  solutions  are  added  to  fibrinogen  solutions 
fibrin  is  formed.  By  Morawitz'  view  the  appearance  of  thrombin  in  the 
blood  is  due  to  the  interaction  of  at  least  three  antecedent  substances. 
These  are,  i,  prothrombin  (thrombogen),  2,  calcium,  and  3,  thrombo- 
kinase  (cytozym).  If  the  blood  is  drawn  from  vessels  with  due  pre- 
cautions, i.e.,  not  to  allow  it  to  come  in  contact  with  the  cut  vessel  or  other 
tissue,  clotting  is  very  much  delayed.  The  plasma  if  separated  by  the 
centrifuge  will  remain  unclotted  for  a  long  time  as  shown  by  Howell  for 
the  terrapin's  plasma.  This  plasma  will  quickly  clot  at  any  time  if  a  few 
drops  of  tissue  extract  in  salt  solution  be  added.  When  blood  is  drawn  over 
the  lacerated  tissue  of  a  wound  it  clots  more  quickly.  These  observations 
have  led  to  the  assumption  of  an  activating  substance  or  kinase  to  which 
the  name  thrombokinase  has  been  given  by  Morawitz.  It  is  assumed  to 
have  its  origin  in  tissue  cells  and  in  the  cells  of  the  blood,  especially  the 
leucocytes. 

If  precautions  are  taken  to  draw  the  blood  in  such  a  manner  as  to  re- 
move the  calcium  from  the  plasma,  no  clot  is  formed.  The  calcium  which 
exists  in  solution  in  the  plasma  to  the  extent  of  0.026  per  cent,  (measured 
as  calcium  chloride)  can  be  removed  by  precipitation  with  oxalate  solution 
or  by  the  action  of  fluorides  or  citrates  which  bind  the  calcium  so  that  it  is 
no  longer  available  to  the  prothrombin.  Such  plasma  contains  fibrinogen, 
prothrombin,  and  thrombokinase,  and  whenever  calcium  chloride  is  added 
to  excess,  coagulation  takes  place. 

L.  J.  Rettger  has  recently  made  a  re-examination  of  the  conditions  for  the 
clotting  of  blood.  He  questions  the  interpretation  of  the  facts  on  the  basis  of 
which  the  assumption  of  a  kinase  is  made.  He  says: 

"Terrapin's  blood  taken  carefully  through  an  oiled  cannula  and  put  into 
a  perfectly  clean  beaker  will  remain  fluid  for  days.  The  blood  may  be  cen- 
trifuged,  and  the  clear  supernatant  plasma  is  then  equally  or  more  resistant 
to  spontaneous  clotting.  If,  however,  tissue  extracts  or  pieces  of  tissue  be 
added,  the  coagulation  is  pronounced  and  immediate.  The  most  apparent 
explanation  is  that  a  thrombin  or  coagulin  or  kinase  is  present  in  the  extract. 
But  this  blood  or  plasma  can  be  made  to  clot  equally  well  and  equally  rapidly 
in  ways  which  preclude  the  presence  of  such  a  definite  agent.  If,  for  instance, 


THEORIES  OF  COAGULATION  123 

dust  particles,  loose  sweepings,  or  linty  shreds  be  added,  the  coagulation  is 
equally  prompt  and  in  a  number  of  experiments  was  more  rapid  than  with  tis- 
sue extract."  "The  bird's  blood  or  plasma  clots  with  practically  the  same 
rapidity  and  firmness  if  dust  particles  are  generously  added.  Bits  of  down 
or  feathers  introduced  bring  about  a  speedy  clotting.  Surely  there  can  be 
no  question  of  a  'kinase'  in  these  instances."  "The  existence  of  'kinase' 
or  'coagulins'  in  the  various  tissues  is  improbable.  Using  carefully  pre- 
pared fibrinogen  solutions,  extracts  of  tissues,  irrigated  to  remove  every 
trace  of  thrombin-containing  blood,  cause  no  clotting.  When  the  addition 
of  such  extracts  produces  coagulation  in  bloods  of  bird  or  reptile,  similar 
results  can  be  secured  by  the  addition  of  substances,  such  as  dust,  lint, 
shreds,  etc.,  which  preclude  the  presence  of  definite  coagulating  agents. 
These  results  render  very  probable  the  assumption  that  in  such  plasmas  all 
the  factors  of  coagulation  are  in  reality  present,  and  the  addition  of  tissue 
extract  or  other  foreign  substance  brings  into  the  mixture  nothing  that  can 
be  regarded  as  a  coagulin  or  as  a  kinase." 

Howell  has  been  studying  the  phenomena  of  coagulation  for- a  number  of 
years.  On  the  basis  of  his  work  he  makes  a  somewhat  different  interpreta- 
tion of  the  facts  on  which  the  theory  of  Morawitz  is  founded.  By  Howell's 
view,  "Circulating  blood  contains  normally  all  the  necessary  fibrin  factors, 
namely,  fibrinogen,  prothrombin,  and  calcium.  These  substances  are  pre- 
vented from  reacting,  and  the  normal  fluidity  of  the  blood  is  maintained,  by 
the  fact  that  antithrombin  is  also  present,  and  this  substance  prevents  the 
calcium  from  activating  the  prothrombin  to  thrombin.  In  shed  blood  the 
restraining  effect  of  the  antithrombin  is  neutralized  by  the  action  of  a  sub- 
stance (thromboplastin) ,  furnished  by  the  tissue  elements.  In  the  mammalia 
the  thromboplastin  is  derived,  in  the  first  place,  from  the  elements  of  the 
blood  itself  (blood  platelets).  In  the  lower  vertebrates  the  supply  of  this 
material,  in  normal  clotting,  comes  from  the  external  tissues."  Howell's 
thromboplastin  and  Morawitz'  thrombokinase  are  probably  one  and  the 
same  substance,  it  being  an  enzyme  by  Morawitz'  view,  a  property  denied 
by  Howell  and  by  Rettger. 

Quite  recently  Howell  has  thrown  light  on  the  nature  of  his  thromboplastic 
substance.  He  finds  that  the  lipoid,  kephalin,  present  in  many  tissues  of  the 
body  possesses  the  power  of  neutralizing  antithrombin  and  is  comparable,  in 
relation  to  blood  clotting,  to  the  action  of  thromboplastin.  Lecithin  does  not 
possess  this  property. 

One  may  restate  Morawitz'  view  in  a  word  as  follows:  The  coagulation  of 
the  blood  takes  place  because  of  the  formation  of  fibrin  from  fibrinogen  by 
the  action  of  thrombin.  The  fibrinogen  is  constantly  present  in  the  plasma. 
The  thrombin  is  formed  by  the  interaction  of  three  substances,  prothrombin, 
calcium,  and  thrombokinase.  The  prothrombin  arises  chiefly  from  the  dis- 
integration of  the  blood  platelets  and  leucocytes  when  the  blood  leaves  the 


124  THE    BLOOD 

blood  vessels.  The  calcium  is  present  in  the  blood  plasma  at  all  times. 
The  thrombokinase  originates  in  tissue  cells  of  the  blood  and  of  the  organs  of 
the  body  in  general. 

Rettger's  view  is  best  given  in  his  own  words:  "  On  the  basis  of  the  work 
here  presented  it  is  not  necessary  to  assume  the  existence  of  a  kinase  in  explain- 
ing the  clotting  of  shed  blood.  The  prothrombin  formed  from  the  platelets 
and  leucocytes  by  secretion  or  by  processes  of  disintegration  is  activated  to 
thrombin  by  the  calcium  salts  present,  and  the  thrombin  so  formed  combines 
in  quantitative  fashion  with  the  fibrinogen  to  form  fibrin." 

Howell's  demonstration  of  antithrombin  offers  a  new  factor  in  the 
problem  of  blood  clotting.  For  example,  the  hastening  influence  of  keph- 
alin  on  blood  clotting  is  probably  due  to  its  action  on  antithrombin, 
rather  than  on  either  of  the  other  clotting  complexes. 

The  Coagulation  Time  of  Blood. — The  rapidity  with  which  blood  co- 
agulates varies  greatly  in  different  animals.  In  the  majority  of  mammals 
the  coagulation  time  varies  from  2.5  to  5  minutes.  In  man  this  time  is 
about  3  to  3:5  minutes  in  normal  blood.  In  recent  experiments  by  Cannon 
and  Mendenhall,  on  the  coagulation  time  of  the  blood  from  the  dog  and 
cat,  determined  by  a  new  and  mechanically  accurate  method,  the  normal 
coagulation  time  is  given  as  from  3 . 5  to  4. 5  minutes.  This  coagulation  time 
however  varies  under  different  conditions  of  the  animal,  especially  under 
conditions  which  affect  the  activities  of  the  glands,  in  particular  the  epine- 
phros.  Stimulation  of  this  gland  either  directly  by  stimulation  of  the 
splanchnic  nerves,  or  indirectly  through  conditions  that  arouse  fear,  etc., 
leads  to  a  sharp  diminution  in  the  coagulation  time  of  the  blood.  This 
decrease  in  some  instances  is  well  within  i  minute,  less  than  ^  minute  in 
their  experiment  3,  which  was  an  experiment  after  emotional  excitement. 
The  coagulation  time  is  retarded  by  the  elimination  of  the  circulation  of 
the  intestine  and  of  the  liver,  also  by  nephrectomy. 

Conditions  Affecting  Coagulation. — From  the  preceding  discussion 
it  is  evident  that  the  rapidity  of  the  coagulation  of  the  blood  will  be 
influenced  by  anything  that  will  influence  the  formation  of  the  fibrin 
factors  or  their  interaction.  The  most  important  influences  are  the 
following: 

Condition  of  the  Blood. — The  blood  varies  greatly  through  a  wide  range 
in  its  ability  to  form  fibrin.  This  depends  upon  the  interaction  of  those 
tissues  that  produce  the  fibrin  factors.  An  efficient  circulation  through  the 
abdominal  viscera  is  necessary  to  maintain  the  clotting  properties  of  the 
blood.  The  rapidity  of  clotting  is  increased  following  the  process  of 
digestion.  It  is  also  increased  (Cannon)  by  the  injection  of  epinephrin  or 
by  the  stimulation  of  the  splanchnic  nerves  which  increase  the  output  of 
epinephrin  by  the  suprarenal  bodies.  (See  Influence  of  the  Ductless 
Glands,  page  482).  If  those  glands  be  removed,  the  time  of  blood  clotting 


CONDITION    OF    THE   BLOOD 


125 


increases.  The  exclusion  of  the  abdominal  circulation  tends  to  increase 
the  time  of  blood  clotting,  and  if  the  liver  circulation  be  eliminated,  the 
influence  of  epinephrin  is  lost. 

Hemorrhage  increases  the  rapidity  of  coagulation,  apparently  by  stimu- 
lating the  production  of  the  fibrin  factors. 


FIG.  1 08. — Diagram  of  the  Graphic  Coagulometer.  C, 
In  a  cannula  of  convenient  working  size,  2  mm.  internal 
diameter,  and  2  cm.  long,  which  when  filled  with  a  sample 
of  fresh  circulating  blood  is  quickly  plugged  with  wax  and 
connected  by  a  short  rubber  tube  with  a  longer  glass  tube 
held  in  the  support  U.  The  whole  is  then  immersed  in 
a  beaker  of  water  at  constant  temperature  (not  shown  in  • 
the  figure).  D,  A  copper  wire  of  standard  weight  hanging 
over  the  lever,  the  looped  end  is  immersed  in  the  sample  of 
blood,  C.  W,  Counterpoising  weight  for  the  lever  which 
rotates  on  the  axis  A.  S,  Lever  support.  RX-R2,  Short  L-shaped  arm  which  when 
moved  from  P1  to  P2  releases  the  lever  at  R1.  To  use  the  instrument,  draw  the 
sample  of  blood  at  a  signal,  insert  it  in  the  apparatus,  and  at  regular  intervals  release 
the  lever  by  moving  the  arm  R2.  If  the  blood  is  fluid,  the  counterpoising  lever  will 
make  a  vertical  stroke  through  its  free  range.  If  threads  of  fibrin  have  formed,  these 
counteract  the  movement  of  the  lever.  E,  Time  signal.  From  Cannon. 

Condition  of  the  Blood-vessel  Walls. — Intravascular  clotting  often  takes 
place  upon  injury  of  the  endothelial  lining  of  the  blood  vessels,  either  from 
the  liberation  of  thrombokinase  in  quantity  too  great  for  elimination  by  the 
healthy  portion  of  the  wall  (Morawitz),  or  by  the  disturbance  of  the  equi- 
librium of  the  forces  which  prevent  the  interaction  of  the  fibrin  factors 
present  in  the  blood  (Rettger).  The  healthy  endothelium  no  doubt  is  an  im- 
portant factor  in  controlling  the  relative  amounts  of  the  fibrin  factors  that 
must  be  constantly  forming.  The  open  wounds  and  lacerations  of  tissue 
that  accompany  the  loss  of  blood  by  accident  are  the  very  conditions  most 
favorable  to  clotting,  since  large  amounts  of  tissue  extract  are  set  free  under 
these  conditions. 

Temperature. — Cold  retards  coagulation.  Gentle  warmth,  40°  C.,  has- 
tens, but  a  temperature  above  56°  C.  destroys  clotting,  since  that  temperature 
heat-coagulates  the  fibrinogen. 

Contact  "with  Foreign  Bodies. — Such  contact  hastens  clotting.  This  is 
due  to  the  influence  of  such  bodies  in  hastening  the  formation  of  fibrin 
factors,  especially  the  substances  that  arise  from  the  disintegration  of 
leucocytes. 

Neutral  Salts. — The  additon  of  neutral  salts  in  the  proportion  of  2  or  3 
per  cent,  and  upward  delays  coagulation.  When  added  in  large  propor- 
tions, most  of  these  saline  substances  prevent  coagulation  altogether. 
Coagulation,  however,  ensues  on  dilution  with  water.  The  time  during 
which  salted  blood  can  be  thus  preserved  in  a  liquid  state,  and  coagulated 


126  THE    BLOOD 

by  the  addition  of  water,  is  quite  indefinite,  if  measures  be  taken  to  pre- 
vent putrefaction. 

Oxalates  and  Fluorides. — Oxalates  to  the  extent  of  o.i  per  cent,  con- 
centration prevent  clotting  by  the  removal  of  calcium,  one  of  the  factors 
in  the  formation  of  thrombin.  Once  thrombin  is  formed,  clotting  will 
take  place  in  the  absence  of  soluble  calcium.  This  is  proven  by  the  fact 
that  solutions  of  pure  fibrinogen  and  thrombin  form  fibrin  clots. 

Flourides,  on  the  other  hand,  not  only  precipitate  soluble  calcium  but 
fix  the  blood  platelets  from  which  the  prothrombin  is  formed. 

Peptone. — The  injection  of  commercial  peptone  (a  mixture  of  proteoses 
and  peptones)  in  the  blood  vessels  of  an  animal  to  the  extent  of  o .  5  gram 
of  peptone  per  kilo  weight  of  the  body  of  the  animal  will  deprive  the  blood 
of  the  power  of  coagulation.  If  a  smaller  quantity  be  injected  the  coagula- 
tion of  the  blood  will  be  delayed.  If  peptone  blood  is  drawn  and  centri- 
fuged,  the  plasma  obtained  is  called  peptone  plasma.  Howell  has  shown  that 
peptone  contains  antithrombin  in  a  relatively  large  quantity.  The  increase 
of  antithrombin  acts  to  restrain  the  reaction  of  prothrombin  in  the  formation 
of  thrombin.  Peptone  plasma  in  the  blood  vessels  of  the  animal  gradually 
regains  the  power  to  coagulate.  When  blood  is  drawn  into  a  physiological 
salt  solution  of  proteose-peptone  clotting  occurs. 

MORPHOLOGY  OF  THE  BLOOD. 

The  corpuscles  floating  in  the  fluid  plasma  of  the  blood,  when  separated 
by  a  centrifugal  machine  are  found  to  make  up  45  to  50  per  cent,  of  the  total 
mass  of  the  blood.  These  corpuscles,  or  formed  elements,  are  of  three 
varieties,  the  red  corpuscles  or  erythrocytes,  the  white  corpuscles  leucocytes, 
and  the  blood  platelets  which  have  been  called  thrombocytes. 

Red  Corpuscles  or  Erythrocytes. — Human  red  blood  corpuscles  are 
circular,  biconcave  discs  with  rounded  edges,  from  •j/j.  to  8/*  in  diameter, 
and  about  2/z  in  thickness.  When  viewed  singly  they  appear  of  a  pale 
yellowish  tinge;  the  deep  red  color  which  they  give  to  the  blood  being  ob- 
servable in  them  only  when  they  are  seen  en  masse.  They  are  composed 
of  a  colorless,  structureless,  and  transparent  filmy  framework  or  stroma, 
infiltrated  in  all  parts  by  the  red  coloring  matter,  the  hemoglobin.  The 
stroma  is  tough  and  elastic,  so  that  as  the  corpuscles  circulate  they  admit 
of  elongation  and  other  changes  of  form  in  adaption  to  the  vessels,  yet 
recover  their  natural  shape  as  soon  as  they  escape  from  compression. 

Number  and  Character  of  the  Red  Corpuscles. — The  normal  number  of 
red  blood-cells  in  a  cubic  millimeter  of  human  blood  was  estimated  by 
Welcker,  in  1854,  to  be  5,000,000  in  men  and  4,500,000  in  women.  Num- 
erous recent  observations,  however,  have  shown  that  these  estimates  are  a 
little  low,  especially  in  men,  and  the  average  number  has  been  placed  by 
different  authorities  at  various  points  between  5,000,000  and  5,500,000. 
Still  the  original  numbers  as  given  by  Welcker  are  accepted  at  the  present 


MORPHOLOGY    OF    THE    BLOOD 


127 


day  as  being  sufficiently  accurate  for  ordinary  purposes.  It  has  been  also 
shown  that  there  are  many  distinct  physiological  variations  in  the  number, 
depending  on  the  time  of  day,  digestion,  sex,  etc.  The  number  of  red  cells 
usually  diminishes  in  the  course  of  each  day,  while  the  leucocytes  increase 
in  number.  It  has  been  suggested  that  this  is  due  to  the  influence  of 
digestion  and  of  exercise. 

It  has  generally  been  found  that  within  half  an  hour  or  an  hour  after  a 
full  meal  the  number  of  red  cells  begins  to  diminish,  and  that  this  keeps  up 
for  from  two  to  four  hours,  when  it  is  followed  by  a  gradual  rise  to  the  normal. 
The  usual  fall  is  250,000  to  750,000  per  cubic  millimeter.  These  results 
are  most  marked  after  a  largely  fluid  meal,  and  are  probably  due  to  dilution 
of  the  blood  as  a  result  of  the  absorption  of  fluids.  In  animals  the  number 
of  red  cells  is  increased  by  fasting,  but  in  man  the  results  are  variable,  some 


FIG.  109. 


FIG.  no. 
The  rounded  or  uncolored  corpuscles  are 


FIG.  109. — Red  Corpuscles  in  Rouleaux, 
leucocytes. 

FIG.  no. — Corpuscles  of  the  Frog.  The  central  mass  consists  of  nucleated  colored 
corpuscles.  The  other  corpuscles  are  two  varieties  of  the  colorless  form. 

authorities  claiming  an  increase  and  others  a  decrease.  In  childhood  there 
is  no  difference  between  the  sexes  in  the  number  of  red  cells  per  cubic  milli- 
meter, but  after  menstruation  is  established  a  relative  anemia  develops  in 
women.  Welcker's  original  estimate  placed  the  difference  at  500,000  per 
cubic  millimeter,  and  these  figures  have  been  generally  accepted,  though 
Leichtenstein  asserts  that  the  difference  is  1,000,000. 

Menstruation  in  healthy  subjects  has  practically  no  effect,  as  not  more 
than  100-200  cubic  centimeters  of  blood  are  lost  normally  in  the  course  of 
several  days.  Under  such  circumstances  the  normal  diminution  of  red  cells 
per  cubic  millimeter  is  probably  less  than  150,000,  though  Sfameni  has  placed 
the  loss  at  about  225,000.  The  leucocytes  are  slightly  increased  during 
menstruation.  It  is  now  the  general  opinion  that  pregnancy  has  little  or  no 
effect  on  the  number  of  red  cells,  and  that  any  anemia  must  be  due  to  abnormal 
conditions.  Post-partum  anemia  should  not  last  longer  than  two  weeks. 


128 


THE    BLOOD 


The  red  corpuscles  are  not  all  alike.  In  almost  every  specimen  of  blood  a 
certain  number  of  corpuscles  smaller  than  the  rest  may  be  observed.  They 
are  termed  microcytes,  or  hematoblasts,  and  are  probably  immature  corpuscles. 

A  peculiar  property  of  the  red  corpuscles,  which  is  exaggerated  in  in- 
flammatory blood,  may  be  here  again  noticed,  i.  e,,  their  great  tendency  to 


FIG.  in. — Illustration  exhibiting  the  typical  characters  of  the  red  blood-cells  in  the 
main  Divisions  of  the  Vertebrata.  The  fractions  are  those  of  an  inch,  and  represent  the 
average  diameter.  In  the  case  of  the  oval  cells,  only  the  long  diameter  is  here  given.  It  is 
remarkable,  that  although  the  size  of  the  red  blood-cells  varies  so  much  in  the  different 
classes  of  the  vertebrate  kingdom,  that  of  the  white  corpuscles  remains  comparatively 
uniform,  and  thus  they  are,  in  some  animals,  much  greater,  in  others  much  less,  than  the 
red  corpuscles  existing  side  by  side  with  them.  Modified  from  Gulliver. 

adhere  together  in  rolls  or  columns  (rouleaux),  like  piles  of  coins.  These 
rolls  quickly  fasten  together  by  their  ends,  and  cluster;  so  that,  when  the 
blood  is  spread  out  thinly  on  a  glass,  they  form  a  kind  of  irregular  network, 
with  crowds  of  corpuscles  at  the  several  points  corresponding  with  the  knots 
of  the  net,  figure  109.  Hence  the  clot  formed  in  such  a  thin  layer  of  blood 
looks  mottled  with  blotches  of  pink  upon  a  white  ground. 


DEVELOPMENT  OF  THE  RED  BLOOD  CORPUSCLES 


129 


The  red  corpuscles  are  constantly  undergoing  disintegration  in  different 
parts  of  the  circulatory  system,  particularly  in  the  spleen.  The  liberated 
hemoglobin  contributes  to  the  formation  of  the  bile  pigments  in  the  liver. 

Development  of  the  Red  Blood  Corpuscles. — The  first  formed 
blood  corpuscles  of  the  human  embryo  differ  much  in  their  general  characters 
from  those  which  belong  to  the  later  periods  of  intra-uterine,  and  to  all 
periods  of  extra-uterine  life.  Their  manner  of  origin  is  at  first  very  simple. 

Surrounding  the  early  embryo  is  a  circular  area,  called  the  vascular  area 
in  which  the  first  rudiments  of  the  blood  vessels  and  blood  corpuscles  are 
developed.  Here  the  nucleated  embryonal  cells  of  the  mesoblast,  from 
which  the  blood  vessels  and  corpuscles  are  to  be  formed,  send  out  processes 
in  various  directions, 
and  these,  joining  to- 
gether, form  an  irregu- 
lar mesh-work.  The 
nuclei  increase  in  num- 
ber, and  collect  chiefly 
in  the  larger  masses  of 
protoplasm,  but  partly 
also  in  the  processes. 
It  appears  that  hemo- 
globin then  makes  its 
appearance  in  certain 
of  these  nucleated  em-  FIG.  112. — Part  of  the  Network  of  Developing  Blood 

brvonal       cells      which     Vessels  in  the  Vascular  Area  of  a  Guinea-pig.    Showing  blood 
corpuscles  becoming  free  in  an  enlarged  and  hollowed-out 

thus  become  the  earliest  part  of  the  network  and  processes  of  protoplasm.  (E.  A. 
red  blood  corpuscles.  Schafer.) 

The  protoplasm  of  the  cells  and  their  branched  network  in  which  these, 
corpuscles  lie  then  become  hollowed  out  into  a  system  of  canals  enclosing 
fluid  in  which  the  red  nucleated  corpuscles  float.  The  corpuscles  at  first 
are  from  about  IOJJL  to  i6/<  in  diameter,  mostly  spherical,  and  with  granular 
contents,  and  a  well-marked  nucleus.  Their  nuclei,  which  are  about  5^ 
in  diameter,  are  central,  circular,  very  little  prominent  on  the  surfaces  of 
the  corpuscles,  and  apparently  slightly  granular. 

The  corpuscles  then  strongly  resemble  the  colorless  corpuscles  of  the 
fully  developed  blood  but  for  their  color.  They  are  capable  of  ameboid 
movement  and  multiply  by  division. 

When,  in  the  progress  of  embryonic  development,  the  liver  is  formed, 
the  multiplication  of  blood-cells  in  the  whole  mass  of  blood  ceases,  and  new 
blood-cells  are  produced  by  this  organ,  and  also  by  the  spleen.  These  are 
at  first  colorless  and  nucleated,  but  afterward  acquire  the  ordinary  blood 
tinge,  and  resemble  very  much  those  of  the  first  set.  They  also  multiply  by 
division.  The  bone  marrow  also  begins  to  form  red  corpuscles,  though  at 


130  THE    BLOOD 

first  in  small  amounts  only.  This  function  develops  rapidly,  however,  so 
that  at  birth  the  marrow  represents  the  chief  seat  of  production  of  the  red 
cells.  Nevertheless,  nucleated  red  cells  are  usually  found  at  birth,  sometimes 
in  considerable  quantities  in  the  liver  and  in  the  spleen.  Non-nucleated 
red  cells  begin  to  appear  soon  after  the  first  month  of  fetal  life,  and  gradually 
increase  so  that  at  the  fourth  month  they  form  one-fourth  of  the  whole  amount 
of  colored  corpuscles.  At  the  end  of  fetal  life  they  almost  completely  re- 
place the  nucleated  cells.  In  late  fetal  life  the  red  cells  are  formed  in  almost 
the  same  way  as  in  extra-uterine  life. 


FIG.  113.  FIG.  114. 

FIG.  113. — Multiplication  of  the  Nucleated  Red  Corpuscles.  Marrow  of  young 
kitten  after  bleeding,  showing  above  karyokinetic  division  of  erythroblast,  and  below  the 
formation  of  mature  from  immature  erythrocytes.  (Howell.) 

FIG.  114. — Shows  the  Way  in  which  the  Nucleus  Escapes  from  the  Nucleated  Red 
Corpuscles,  i,  2,  3,  4,  represent  different  stages  of  the  extrusion  noticed  upon  the  living 
corpuscles,  a,  Specimen  from  the  circulating  blood  of  an  adult  cat,  bled  four  times;  b, 
specimen  from  the  circulating  blood  of  a  kitten  forty  days  old,  bled  twice;  c,  specimens 
from  the  blood  of  a  fetal  cat,  9  cm.  long.  Others  from  the  marrow  of  an  adult  cat,  two  of 
the  figures  showing  the  granules  present  in  the  corpuscles,  which  have  been  interpreted 
erroneously  as  a  sign  of  the  disintegration  of  the  nucleus.  (Howell.) 

Various  theories  have  prevailed  as  to  the  mode  of  origin  of  the  non- 
nucleated  colored  corpuscles.  For  a  time  it  was  thought  that  they  were  of 
endoglobular  origin,  and  merely  fragments  of  some  original  cell,  being  pro- 
duced by  subdivision  of  the  cell  body  itself.  This  theory  easily  accounted 
for  the  absence  of  the  nuclei,  but  it  has  not  been  supported  by  recent  investi- 
gations. At  present  it  is  the  general  belief  that  the  non-nucleated  cells,  or 
erythrocytes,  are  derived  from  nucleated  cells  by  a  process  of  mitotic  division, 
and  further  that  their  nuclei  gradually  shrink  or  fade  and  are  then  extruded. 
The  use  of  some  of  the  more  recent  stains  seems  to  prove  that  there  are  traces 
of  nuclear  material  in  the  non-nucleated  corpuscles. 

After  infancy  and  early  childhood  the  origin  of  erythrocytes  is  practically 
limited  to  the  red  marrow  of  the  bones.  The  mother  cells,  or  erythroblasts, 
are  constantly  forming  and  setting  free  erythrocytes,  the  rate  varying  greatly 
at  different  periods. 


THE  COLORLESS  CORPUSCLES  OR  LEUCOCYTES         131 

The  Colorless  Corpuscles  or  Leucocytes.— In  human  blood  the  white 
corpuscles,  leucocytes,  are  nearly  spherical  masses  of  granular  protoplasm 
without  cell  wall.  In  all  cases  one  or  more  nuclei  exist  in  each  corpuscle. 
The  corpuscles  vary  considerably  in  size,  but  average  IO/JL  in  diameter. 

The  number  of  leucocytes  in  a  cubic  millimeter  of  blood  is  estimated 
at  7,500  to  8,000.  The  proportion  of  white  corpuscles  to  red,  therefore,  is 
about  one  of  the  former  to  700  of  the  latter.  This  proportion  is  not  very 
constant  in  health  and  great  variations  occur  under  the  influence  of  disease 


FIG.    115. — Colored  Nucleated  Corpuscles,  from  the  Red  Marrow  of  the   Guinea-pig. 

(E.  A.  Schafer.) 


especially  in  certain  infectious  diseases  in  which  the  number  of  white  cor- 
puscles is  markedly  increased. 

After  a  full  meal  the  white  cells  in  a  healthy  adult  are  increased  in  number 
about  one-third,  the  increase  beginning  within  an  hour,  attaining  a  maxi- 
mum in  three  or  four  hours,  and  then  gradually  falling  to  normal.  This 
process  is  frequently  modified  by  the  character  of  the  food,  the  greatest 
increase  occurring  with  an  exclusively  meat  diet,  while  a  purely  vegetarian 
diet  has  usually  no  effect.  The  increase  is  also  more  marked  in  children, 
and  especially  in  infants.  The  essential  factor  is  probably  the  absorption 
of  albuminous  matter  in  considerable  quantities.  This  causes  proliferation 
of  leucocytes  in  the  lymphoid  tissue  of  the  gastro-intestinal  tract. 

In  pregnancy  there  is  often  a  moderate  increase  in  the  number  of  white 
cells  during  the  later  months.  This  does  not  begin  until  after  the  third 
month,  and  is  most  marked  and  constant  in  primiparae.  After  parturition 
the  leucocytes  gradually  diminish  under  normal  conditions,  and  usually 
reach  the  normal  within  a  fortnight.  The  essential  factor  is  probably  the 
general  stimulation  in  the  maternal  organism.  It  is  well  established  that  the 
white  cells  are  very  numerous  in  the  new-born,  though  different  observers 
have  made  very  conflicting  estimates.  Still  all  agree  that  there  is  a  very 
rapid  decrease  in  their  numbers  during  the  first  few  days,  and  that  this  is 
followed  by  a  less  marked  increase,  which  continues  for  many  months. 
According  to  Rieder  there  are  at  birth  from  14,200  to  27,400  per  cubic  milli- 
meter, and  after  the  fourth  day  from  12,400  to  14,800. 

The  colorless  corpuscles  present  a  great  diversity  of  form.  There  are 
certain  constant  types  found  in  fairly  definite  proportions  in  normal 
blood,  but  in  pathological  bloods  a  long  series  of  variants  have  been 
described  and  figured  by  such  authors  as  Wood,  Webster,  and  Simon.  In 
histological  and  clinical  examination  the  white  corpuscles  are  classified 


132  THE   BLOOD 

according  to  their  size,  structure,  and  staining  reaction.  Some  of  these 
cells  are  mononuclear,  others  polynuclear  and  many  charged  with  special 
types  of  granules  that  take  now  basic,  now  acid  dyes,  presumably  accord- 
ing to  their  clinical  composition.  The  following  varieties  may  be  listed 
for  normal  adult  blood. 

1.  Small  mononuclear  leucocytes,  22-25. 

2.  Large  mononuclear  leucocytes,  1-2. 

3.  Polynuclear  neutrophilic  leucocytes,  65-75. 

4.  Polynuclear  eosinophilic  leucocytes,  1-4. 

5.  Polynuclear  basophilic  leucocytes,  mast  cells,  0.2  to  0.5. 

The  small  mononuclear  leucocytes,  or  lymphocytes,  are  about  the  size 
of  or  smaller  than  the  red  corpuscles,  a  single  nucleus  with  very  little 
nongranular  protoplasm,  staining  deeply  with  methyline  blue  with  a 
lighter  staining  nucleus,  22  to  25  per  cent. 

The  large  mononuclear  leucocytes  are  double  the  size  of  the  small  leuco- 
cytes or  even  larger.  They  have  a  single  nucleus  about  the  size  of  the 
preceding  type  but  a  much  larger  relative  development  of  protoplasm. 
Their  cytoplasm  is  not  granular  and  they  are  weakly  basophilic.  These 
cells  like  the  small  leucocytes  arise  in  the  lymphoid  tissue. 

The  polynuclear  neutrophilic  leucocytes  are  about  the  size  of  a  red  blood 
corpuscle  and  are  granular  in  appearance.  The  nuclei  take  basic  dyes. 
The  cytoplasm  is  slightly  acidic  but  the  granules  imbedded  in  it  are  baso- 
philic. These  leucocytes  constitute  from  65  to  75  per  cent,  of  the  total 
number  of  white  corpuscles.  This  class  is  most  actively  phagocytic  and 
is  increased  in  number  in  response  to  most  infections. 

The  polynuclear  eosinophilic  leucocytes. — The  cells  of  this  type  are  the 
largest  cells  of  the  white  corpuscle  group.  Their  cytoplasm  is  crowded  with 
granules  which  stain  deeply  with  eosin  and  other  acid  dyes.  From  this 
characteristic  they  get  their  name.  The  eosinophiles  constitute  i  to  4 
per  cent,  of  the  total.  They  are  extremely  motile  and  phagocytic  and  are 
very  greatly  increased  in  number  in  certain  diseases. 

The  mast  cells  are  much  fewer  in  number  except  in  certain  particular 
diseases.  They  have  a  polymorphic  nucleus  characteristic  in  appearance. 
They  take  basic  dyes  with  difficulty.  These  cells  are  also  granular.  The 
granules  do  not  take  eosin  but  are  basic  in  type.  They  vary  in  size,  being 
rather  larger  than  eosinophilic  granules.  In  normal  blood  one  mast  cell 
occurs  in  from  200  to  250  leucocytes. 

The  relative  number  of  leucocytes  varies  in  children  as  compared  with 
normal  adults.  The  small  mononuclear  lymphocytes  are  practically 
double  the  adult  number  and  the  polymorphonuclears  about  half  the 
number  in  the  adult.  The  eosinophiles  are  also  more  frequent  though 
still  relatively  rare. 


THE   BLOOD    PLATES 


133 


FIG.  116— (a)  Red  blood  corpuscle  for  comparison;  (b)  Small  lymphocyte;  (c) 
Large  lymphocyte  (myelocyte);  (d)  Fine  and  (e)  coarse  eosinophiles;  (/)  Basophile. 
(F.  C.  Busch.) 

Ameboid  Movement  and  Phagocytic  Action  of  Leucocytes. — The  remark- 
able property  of  the  colorless  corpuscles  of  spontaneously  changing  their 
shape  was  first  demonstrated  by 
Wharton  Jones  in  the  blood  of  the  skate. 
If  a  drop  of  blood  be  examined  with 
a  high  power  of  the  microscope,  under 
conditions  by  which  loss  of  moisture  is 
prevented,  and  at  the  same  time  the 
temperature  is  maintained  at  about  that 
of  the  body,  37°  C.,  the  colorless  corpus- 
cles will  be  observed  slowly  to  alter 
their  shapes,  and  to  send  out  processes 
at  various  parts  of  their  circumference. 
The  ameboid  movement  which  can  be 

demonstrated  in  human  colorless  blood  FlG  Il8._Blood  Plates>  showing 
Corpuscles  can  be  most  conveniently  chromatic  centers  regarded  by  some  as 
studied  in  the  newt's  blood.  Processes 
are  sent  out  from  the  corpuscle.  These 
may  be  withdrawn,  but  more  often  the  protoplasm  of  the  whole  corpuscle 
flows  gradually  forward  to  the  position  occupied  by  the  process,  thus  the 
corpuscle  changes  its  position.  The  change  of  position  of  the  corpuscle 
can  also  take  place  by  a  flowing  movement  of  the  whole  mass,  and  in 
this  case  the  locomotion  is  comparatively  rapid.  The  activity  both  in 
the  processes  of  change  of  shape  and  also  of  change  in  position  is  much 
more  marked  in  some  corpuscles  than  in  others.  Klein  states  that  in  the 


134 


THE   BLOOD 


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THE    SERUM  135 

newt's  blood  the  changes  are  especially  noticeable  in  a  variety  of 
colorless  corpuscle,  which  consists  of  a  mass  of  finely  granular  protoplasm 
with  jagged  outline  and  contains  three  or  four  nuclei,  or  in  large  irregular 
masses  of  protoplasm  containing  from  five  to  twenty  nuclei. 

The  blood  leucocytes  are  the  phagocytes  of  the  body.  By  means 
of  their  ameboid  movements  they  surround  or  engulf  foreign  bodies 
including  bacteria.  These  they  digest,  therefore  destroy.  It  is  through 
the  activity  of  the  leucocytes  that  the  body  gains  its  relative  immunity, 
since  these  cells  are  able  to  overcome  to  a  certain  extent  bacterial  invasion. 
The  blood  largely  depends  on  the  polymorphonuclear  leucocytes  for 
phagocytosis,  hence  these  cells  are  found  to  be  increased  in  number  under 
the  stimulus  of  infectious  processes.  The  relative  numbers  of  leucocytes 
in  clinical  blood  counts  have  their  value  in  part  in  this  fact.  However,  in 
clinical  bloods  many  pathological  types  appear  which  are  of  special 
significance. 

The  Blood  Platelets  or  Thrombocytes. — A  third  variety  of  corpuscle 
found  in  the  blood  is  known  as  the  blood  platelets.  They  are  circular  or 
elliptical  in  shape,  of  nearly  homogeneous  structure,  and  vary  in  size  from 
0.5  to  5/-1/A.  Hence  they  are  smaller  than  the  red  corpuscles.  They  vary 
in  number  from  5,000  to  45,000  per  cubic  millimeter  and  are  preserved 
by  drawing  fresh  blood  directly  into  Hayem's  or  other  preserving  fluid. 
When  fresh  blood  is  drawn  the  blood  platelets,  coming  in  contact  with 
foreign  bodies,  rapidly  disintegrate  and  give  rise  to  one  of  the  antecedents 
of  blood  plasma  concerned  in  clotting,  prothrombin,  p.  121. 

CHEMICAL  COMPOSITION  OF  THE  BLOOD. 

The  chemical  composition  of  the  blood  as  a  whole  may  be  presented 
by  consideration  of  the  constituents  of  the  plasma  and  of  the  corpuscles. 
The  available  blood  analyses  have  dealt  with  whole  blood,  with  either 
plasma  or  serum  and  with  whole  corpuscles,  although  separate  analyses 
of  red  and  white  corpuscles  have  been  made. 

The  classical  analyses  in  the  literature  that  have  become  the  reliance 
for  teaching  purposes  are  those  of  Schmidt  for  the  blood  of  man  and  of 
Abderhalden  for  dog's  and  horse's  blood.  These  analyses  are  quoted  in 
tabular  form  below. 

ANALYSES   OF  BLOOD   OF  MAN— KARL  SCHMIDT 

One  Thousand  Grammes  of  Blood. 

Blood- Corpuscles 513 . 02 

Water 349 -69 

Substances  not  vaporizing  at  1 20°  163  . 33 

Hematin 7  -7°  (including  0.512  iron) 

Blood  casein,  etc 151 .89 

Inorganic  constituents 3-74  (excluding  iron) 


136 


THE    BLOOD 


Chlorine 0.898 

Sulphuric  acid 0.031 

Phosphoric  acid o .  695 

Potassium i .  586 

Sodium 0.241 

Phosphate  of  lime o .  048 

Phosphate  of  magnesium 0-031 

Oxygen 0.206 

Blood  Plasma 486 . 89 

Water 439 . 02 

Substances  not  vaporizing  at  1 20°     47 . 96 

Fibrin 

Protein,  etc 

Inorganic  constituents 

Chlorine 

Sulphuric 

Phosphoric  acid 

Potassium 

Sodium 

Phosphate  of  lime 

Phosphate  of  magnesium .... 
Oxygen 

Specific  Gravity  =  1.0599. 


Chloride  of  potassium . .  . 
Sulphate  of  potassium . . . 
Phosphate  of  potassium.  . 
Phosphate  of  sodium .... 

Soda 

Phosphate  of  lime 

Phosphate  of  magnesium. 

Total. . 


1.887 
0.068 

1  .  202 
0.325 
0-175 
0.048 

o .  03 1 


3-93 
39.89 

4.14 

1.722 
0.063 
0.071 

0-153 
1.661 

Sulphate  of  potassium  
Chloride  of  potassium  
Chloride  of  sodium  
Phosphate  of  sodium  
Soda 

.  .  .  .   0.137 
.  .  .  .   0.175 

....      2.  701 
0.132 

°  746 

o   14? 

Phosphate  of  lime  

....   o.  145 

*to 
o.  106 

O.  221 

Phosphate  of  magnesium  

....   o  .  i  06 

Total. 


4.142 


The  Composition  of  Plasma. — The  plasma  is  the  liquid  part  of  the 
blood  in  which  the  corpuscles  float.  It  differs  from  the  serum  only  in  that 
the  fibrin  factors  have  been  removed  during  the  process  of  clotting.  To 
all  intents  and  purposes  the  chemistry  of  the  plasma  and  serum  are  iden- 
tical. Plasma  may  be  freed  from  the  corpuscles  by  the  centrifugal 
machine  in  the  interval  before  clotting  forms.  However,  it  is  customary 
to  delay  clotting  by  means  enumerated  below. 

Cooled  Plasma. — If  blood  is  drawn  directly  into  a  chilled  vessel  and 
kept  at  a  temperature  of  about  o°  C.,  or  only  two  or  three  degrees  higher,  the 
corpuscles  will  settle  out,  leaving  a  clear  supernatant  plasma.  This  plasma 
will  clot  promptly  on  raising  the  temperature.  This  method  yields  a 
pure  plasma  for  isolation  of  fibrinogen  and  for  quantitative  analysis. 

Salted  Plasma. — Blood  will  not  clot  if  it  is  mixed  with  sodium  sulphate 
or  magnesium  sulphate  in  adequate  proportion,  one  part  to  twelve  parts 
of  blood  of  the  former,  and  one  part  to  six  parts  of  blood  of  the  latter. 
Salted  plasma  obtained  by  this  method  will  coagulate  on  dilution. 

Oxalated  Plasma. — In  experimental  work  it  is  customary  to  prevent 
blood  clotting  by  receiving  it  into  an  oxalate  solution  or  over  a  powder  of 
an  oxalate  salt.  It  is  necessary  that  the  blood  contain  at  least  one  tenth 
per  cent,  of  oxalate.  The  oxalate  removes  the  soluble  calcium  salts  before 
the  prothrombin  is  converted  into  thrombin.  Oxalated  plasma  will  not 


HEMOGLOBIN  137 

clot  until  a  calcium  salt  is  added.  The  oxalate  does  not  interfere  with  gas 
absorption  and  gas  determination  tests. 

Hirudin  Plasma. — Leach  extract  contains  an  anticoagulant  which 
prevents  clotting  of  blood.  Such  blood  clots  readily  on  adding  serum  or 
other  solution  containing  thrombokinase  which  neutralizes  the  hirudin 
effect  as  it  does  the  antithrombin  when  blood  is  drawn. 

Peptone  Plasma. — Peptone  solution  injected  directly  into  the  blood 
stream  renders  the  blood  non-coaguable  when  it  is  afterward  drawn. 
Peptone  does  not  prevent  blood  from  clotting  when  mixed  in  vitro.  How- 
ever, when  blood  serum  or  a  pure  solution  of  fibrin  ferment  is  added,  both  of 
which  neutralize  the  antithrombin,  then  peptone  plasma  will  form  a  clot. 

Water  of  Blood  Plasma. — The  water  of  the  plasma  varies  in  amount 
through  a  wide  range.  During  absorption  of  food  and  drink  the  water 
increases  temporarily  in  the  blood,  though  at  this  time  it  is  flowing  into 
the  tissues  and  being  more  rapidly  excreted  by  the  kidneys.  In  a  few 
minutes,  say  15  or  20,  after  a  glass  of  water  the  blood  constituents  will  be 
diluted  and  the  red  corpuscles  demonstrably  swollen.  This  condition 
is  quickly  equalized.  On  the  other  hand  the  rapid  loss  of  water  in  evapor- 
ation of  sweat  following  vigorous  exercise  quickly  leads  to  loss  of  water 
from  and  concentration  of  the  plasma.  These  variations  occur  around  an 
average  content  of  90  per  cent,  of  plasma  water. 

Proteins. — The  chief  proteins  of  plasma  are  serum  albumin,  serum 
globulin,  and  fibrinogen,  a  total  of  from  6  to  8  per  cent,  of  the  plasma. 
Fibrinogen  is  the  part  of  plasma  that  is  converted  into  fibrin  when  blood 
clots.  It  is  a  globulin.  Fibrinogen  is  precipitated  from  plasma  by  half 
saturation  with  sodium  chloride  and  along  with  globulin  by  full  saturation 
with  sodium  chloride  or  magnesium  sulphate.  It  is  soluble  in  dilute  salt 
solutions  but  not  soluble  in  water. 

Serum  globulin  or  paraglobulin  is  completely  precipitated  by  saturation 
by  magnesium  sulphate,  incompletely  by  sodium  chloride,  and  coagulates 
at  a  temperature  of  75°  C.  It  is  likewise  soluble  in  dilute  salt  solutions, 
but  insoluble  in  water.  It  is  present  in  plasma  in  from  3.5  to  4  per  cent., 
but  varies  greatly. 

Serum  albumin  is  the  protein  which  predominates  in  human  plasma. 
It  is  readily  obtained  in  crystalline  form ;  is  soluble  in  saturated  magnesium 
sulphate  and  sodium  chloride  solutions,  but  insoluble  in  saturated  ammon- 
ium sulphate  solutions.  It  coagulates  in  neutral  or  acid  solutions  at  from 

73°  to  75°  C. 

Extractives. — The  extractives  are  the  nitrogen-containing  substances, 
such  as  urea,  uric  acid,  creatin,  creatinin,  etc.,  and  the  non-nitrogenous 
glycogen,  dextrose,  cholesterin,  etc.,  a  total  of  0.5  to  0.6  per  cent.  The 
dextrose  content  amounts  to  from  o.i  to  0.15  per  cent. 

Among  the  extractives  must  be  classed  the  various  hydrolytic  ferments 
such  as  the  diastatic  ferments  that  react  with  the  carbohydrates  of  the 


138  THE   BLOOD 

blood,  liver,  etc.;  the  lipolytic  or  fat-splitting  ferments,  and  the  proteolytic 
ferments.  From  their  property  of  producing  not  only  hydrolytic  cleavage 
but  the  reverse  reactions  of  synthesis  these  ferments  hold  one  of  the  most 
significant  positions  among  the  plasma  constituents. 

Inorganic  Substances. — The  total  inorganic  salts  of  human  plasma 
amount  to  from  0.80  to  0.89  per  cent.  By  the  recent  titration  method  of 
Cramer,  the  sodium  ran  323  to  344  milligrams  per  100  cubic  centimeters 
of  serum.  The  chief  salt  is  sodium  chloride  which  constitutes  over  half 
the  total  and  contributes  largely  to  the  osmotic  pressure  of  the  blood. 
Curiously  enough  only  a  minimal  quantity  of  potassium  salt  is  present  in 
the  plasma,  from  18  to  21  milligrams  in  a  hundred  grams.  The  potassium 
exists  as  sulphate  and  basic  phosphate.  The  calcium  is  very  constant  in 
normal  serum,  from  9.3  to  9.9  milligrams  in  a  hundred  cubic  centimeters 
of  serum.  It  is  probably  in  the  blood  as  a  phosphate.  The  calcium  con- 
tent varies  greatly  in  disease,  is  depressed  in  parathyroid  tetany. 

The  Serum. — The  serum  is  the  liquid  part  of  the  blood  or  of  the  plasma 
which  remains  after  the  fibrin  has  been  formed  and  removed.  It  is  a 
transparent  yellowish  fluid  with  a  specific  gravity  of  1025  to  1032.  Serum 
is  ordinarily  obtained  free  from  blood  corpuscles  by  whipping  the  blood, 
that  is  removing  the  fibrin  as  fast  as  it  forms  and  then  sedimenting  the 
corpuscles  in  the  centrifugal  machine.  It  may  be  obtained  by  allowing 
blood  to  clot  in  a  test  tube  or  beaker  and  then  stand  in  the  cold.  The 
clot  contracts,  squeezing  out  the  clear  yellowish  straw-colored  serum.  In 
quantitative  chemical  analysis  the  serum  is  essentially  of  the  same  compo- 
sition as  plasma,  in  fact  serum  is  usually  taken  for  such  quantitative  work. 
It  differs  from  plasma  only  in  the  loss  of  the  fibrin  or  the  fibrin  factors 
which  go  to  form  fibrin.  It  is  usually  rich  in  thrombokinase  or  fibrin 
ferments.  The  percentages  given  above  for  the  salts  of  plasma  were 
actually  determined  on  serum. 

The  Composition  of  the  White  Corpuscles. — The  white  corpuscles 
are  comparatively  undifferentiated  cellular  elements,  hence  possess  the 
chemical  composition  of  protoplasm.  Lillienfeld  has  made  an  analysis 
of  the  leucocytes  of  thymus  gland  from  the  calf,  which  contain  11.49  Per 
cent,  of  solids,  as  follows: 

In  100  Parts  of  Dry  Substance  of  White  Corpuscles  of  Calf. 

Per  cent. 

Protein 1.76 

Leuconuclein 68 .  78 

Histon 8 .  76 

Lecithin 7.51 

Fat 4.02 

Cholesterin 4 .40 

Glycogen o .  80 

96.03 


HEMOGLOBIN 


139 


Most  noteworthy  substances  in  this  table  are  the  nuclein  and  histon, 
first  isolated  by  Kossel  and  Lillienfeld  as  nucleohiston.  Besides  the 
substances  in  the  table,  the  white  corpuscles  contain  salts  of  potassium, 
sodium,  calcium,  and  magnesium,  with  potassium  phosphate  present  in 
greatest  amount. 

The  Composition  of  the  Red  Corpuscles. — Analysis  of  moist  blood 
corpuscles  shows  the  following  results: 

Water 68.8  per  cent. 

Solids- 
Organic 30-388 


Mineral..  0.812  f       3I'2 


Of  the  solids  the  most  important  is  the  respiratory  pigment,  hemoglobin, 
the  substance  to  which  the  blood  owes  its  color.  It  constitutes,  as  will  be 
seen  from  the  appended  table,  more  than  90  per  cent,  of  the  organic  matter 
of  the  corpuscles.  Besides  hemoglobin  the  corpuscles  contain  protein  and 
fatty  matters,  the  former  chiefly  consisting  of  globulins,  and  the  latter  of 
cholesterol  and  lecithin. 

In  100  parts  of  organic  matter  are  found: 

Hemoglobin 90 . 54  per  cent. 

Proteins 8.67  per  cent. 

Fats o .  79  per  cent. 


100.0 


The  inorganic  salts  of  the  red  corpuscles  differ  from  the  salts  of  serum 
in  that  the  ash  of  corpuscles  contains  a  high  content  of  those  salts  that 
tend  to  form  fixed  organic  compounds.  For  example,  iron  is  present  as  a 
part  of  the  hemoglobin  molecule.  There  is  an  excess  of  potassium  in 
corpuscles,  present  in  fixed  organic  compounds.  Only  a  small  amount 
of  sodium  is  present,  and  of  calcium  only  a  trace. 

Hemoglobin. — Of  the  substances  in  the  erythrocytes,  by  far  the  most 
important  from  every  point  of  view  is  the  pigment,  hemoglobin.  It  composes 
about  90  per  cent,  of  the  total  solids  of  the  corpuscles;  therefore,  between 
14  and  15  per  cent,  of  the  blood  itself.  Hemoglobin  is  the  most  complex 
compound  in  the  body,  having  a  molecule  of  the  enormous  molecular  weight 
of  16,669.  Hemoglobin  is  intimately  distributed  throughout  the  stroma  of 
the  corpuscle,  and  when  dissolved  out  it  can  be  crystallized. 

Its  percentage  composition  is  C,  53.85;  H,  7.32;  N,  16.17;  O,  21.84;  8,0.63 
Fe,  0.42.  Jacquet  gives  the  empirical  formula  for  the  hemoglobin  of  the  dog, 
C7S8H12o5N195S3FeO218.  The  most  interesting  of  the  properties  of  hemo- 
globin are  its  powers  of  crystallizing  and  its  attraction  for  oxygen  and  other 
gases  under  certain  pressure  relations. 

Hemoglobin  Crystals. — The  hemoglobin  (oxy hemoglobin)  of  the  blood  of 
various  animals  possesses  the  power  of  crystallizing  to  very  different  ex- 


140 


THE  BLOOD 


tents.  In  some  the  formation  of  crystals  is  almost  spontaneous,  whereas 
in  others  it  takes  place  either  with  great  difficulty  or  not  at  all.  Among 
the  animals  whose  blood  coloring-matter  crystallizes  most  readily  are  the 
guinea-pig,  rat,  squirrel,  and  dog;  and  in  these  cases  to  obtain  crystals  it 
is  generally  sufficient  to  dilute  a  drop  of  recently  drawn  blood  with  water 
and  to  expose  it  for  a  few  minutes  to  the  air.  In  many  instances  other  means 
must  be  adopted;  e.g.,  the  addition  of  alcohol,  ether,  or  chloroform,  rapid 
freezing  and  then  thawing,  the  application  of  an  electric  current,  a  tempera- 


4\ 


FIG.  119. — Crystals  of  Oxyhemoglobin 
— Prismatic,  from  Human  Blood. 


FIG.     1 20. — Oxyhemoglobin     Crystals — 
Tetrahedral,  from  Blood  of  the  Guinea-pig. 


ture  of  60°  C.,  the  addition  of  sodium  sulphate,  or  the  addition  of  decom- 
posing serum  of  another  animal. 

The  hemoglobin  of  human  blood  crystallizes  with  difficulty,  as  does  also 
that  of  the  ox,  the  pig,  the  sheep,  and  the  rabbit. 

The  forms  of  hemoglobin  crystals,  as  will  be  seen  from  figures  119  and 
1 20,  differ  greatly.  Hemoglobin  crystals  are  soluble  in  water.  Both  the 
crystals  themselves  and  also  their  solutions  have  the  characteristic  color  of 
arterial  blood. 

A  dilute  solution  of  Oxyhemoglobin  gives  a  characteristic  appearance 
with  the  spectroscope.  Two  absorption  bands  are  seen  between  the  solar 
lines  D,  which  is  the  sodium  band  in  the  yellow,  and  E,  see  the  frontispiece,, 
one  in  the  yellow,  with  its  middle  line  some  little  way  to  the  right  of  D.  This 
band  is  very  intense,  but  narrower  than  the  other,  which  lies  in  the  green 
near  to  the  left  of  E.  Each  band  is  darkest  in  the  middle  and  fades  away 
at  the  sides.  As  the  strength  of  the  solution  increases,  the  bands  become 
broader  and  deeper.  Both  the  red  and  the  blue  ends  of  the  spectrum  become 
encroached  upon  until  the  bands  coalesce  to  form  one  very  broad  band  when 
only  a  slight  amount  of  the  green  and  part  of  the  red  remain  unabsorbed. 
Any  further  increase  of  strength  leads  to  complete  absorption  of  the  spectrum. 

If  crystals  of  hemoglobin  are  exposed  to  an  atmosphere  of  oxygen  they 
take  up  oxgyen  and  form  Oxyhemoglobin,  each  gram  of  the  pigment  fixing 


HEMOGLOBIN  141 

a  definite  amount  oxygen,  see  chapter  on  Respiration.  When  subjected 
to  a  mercurial  air-pump  the  oxygen  is  given  off,  and  the  crystals  become 
of  a  purple  color.  A  solution  of  the  oxyhemoglobin  in  the  blood  corpuscles 
may  be  made  to  give  up  oxygen,  and  to  change  color  in  a  similar  manner. 
One  gram  of  oxyhemoglobin  liberates  i .  59  c.c.  oxygen,  or,  according  to  Hiif- 
ner's  later  determinations,  i  .34  c.c.,  see  page  292. 

This  change  may  be  also  effected  by  passing  through  the  solution  of 
blood  or  of  oxyhemoglobin,  hydrogen  or  nitrogen  gas,  or  by  the  action  of 


FIG.  121. — Hexagonal  Oxyhemoglobin  Crystals,  from  Blood  of  Squirrel.  On  these 
hexagonal  plates  prismatic  crystals,  grouped  in  a  stellate  manner,  not  unfrequently  occur 
(alter  Funke). 

reducing  agents,  of  Stokes's  fluid*  or  ammonium  sulphide  are  the  most 
convenient. 

With  the  spectroscope,  a  solution  of  deoxidized  or  reduced  hemoglobin 
is  found  to  give  an  entirely  different  appearance  from  that  of  oxidized  hemo- 
globin. Instead  of  the  two  bands  at  D  and  E,  we  find  a  single  broader  but 
fainter  band  occupying  a  position  midway  between  the  two,  and  at  the  same 
time  less  of  the  blue  end  of  the  spectrum  is  absorbed.  Even  in  strong  solu- 
tions this  latter  appearance  is  found,  thereby  differing  from  the  strong  solu- 
tion of  oxidized  hemoglobin  which  lets  through  only  the  red  and  orange 
rays;  accordingly,  to  the  naked  eye  the  one  (reduced-hemoglobin  solution) 
appears  purple,  the  other  (oxyhemoglobin  solution)  red.  The  deoxidized 
crystals  or  their  solutions  quickly  absorb  oxygen  on  exposure  to  the  air, 
becoming  scarlet.  If  solutions  of  blood  be  taken  instead  of  solutions  of 
hemoglobin,  results  similar  to  the  whole  of  the  foregoing  can  be  obtained. 

Venous  blood  never,  except  in  the  last  stages  of  asphyxia,  fails  to  show 
the  oxyhemoglobin  bands,  inasmuch  as  the  greater  part  of  the  hemoglobin 
even  in  venous  blood  exists  in  the  more  highly  oxidized  condition. 

*Stokes's  fluid  consists  of  a  solution  of  ferrous  sulphate,  to  which  ammonia  has  been 
added  and  sufficient  tartaric  acid  to  prevent  precipitation.  Another  reducing  agent  is  a 
solution  of  stannous  chloride,  treated  in  a  way  similar  to  the  ferrous  sulphate,  and  a  third 
reagent  of  like  nature  is  an  aqueous  solution  of  yellow  ammonium  sulphide,  - 


142  THE    BLOOD 

Action  of  Gases  on  Hemoglobin. — Carbon  monoxide  gas  passed  through 
a  solution  of  hemoglobin  causes  it  to  assume  a  cherry-red  color  and  to 
present  a  slightly  altered  spectrum;  two  bands  are  still  visible  but  are 
slightly  nearer  the  blue  end  than  those  of  oxyhemoglobin,  see  Plate  I. 
The  amount  of  carbon  monoxide  taken  up  is  equal  to  the  amount  of  the 
oxygen  displaced.  Carbon  monoxide  gas  readily  displaces  oxygen  under 
the  ordinary  respiratory  conditions.  It  is  less  readily  displaced  by  excess 
of  oxygen  and  by  carbon  dioxide,  hence  the  poisonous  effects  of  coal  gas 
which  contains  much  carbon  monoxide.  Carbon  monoxide  hemoglobin 
is  not  an  oxygen  carrier,  and  death  may  result  from  suffocation  due  to  the 
want  of  oxygen,  notwithstanding  the  free  entry  of  pure  air  into  the  lungs. 
Crystals  of  carbon  monoxide  hemoglobin  closely  resemble  in  form  those 
of  oxyhemoglobin. 

Nitric  oxide  produces  a  similar  compound  to  the  carbon  monoxide 
hemoglobin,  which  is  even  less  easily  reduced. 

Sulphuretted  hydrogen,  if  passed  through  a  solution  of  oxyhemoglobin, 
reduces  it  and  an  additional  band  appears  in  the  red.  If  the  solution  be 
then  shaken  with  air,  the  two  bands  of  oxyhemoglobin  replace  that  of 
reduced  hemoglobin,  but  the  band  in  the  red  persists. 

Methemoglobin. — If  an  aqueous  solution  of  oxyhemoglobin  is  exposed 
to  the  air  for  some  time,  its  spectrum  undergoes  a  change;  the  two  d  and 
e  bands  become  faint  and  a  new  line  in  the  red  at  C  is  developed.  The 
solution,  too,  becomes  brown  and  acid  in  reaction,  and  is  precipitable  by 
basic  lead  acetate.  This  change  is  due  to  the  decomposition  of  oxyhemo- 
globin, and  to  the  production  of  methemoglobin.  On  adding  ammonium 
sulphide,  reduced  hemoglobin  is  produced,  and  on  shaking  this  up  with  air, 
oxyhemoglobin  is  again  produced.  Methemoglobin  is  probably  a  stage  in 
the  deoxidation  of  oxyhemoglobin.  It  appears  to  contain  less  oxygen  than 
oxyhemoglobin,  but  more  than  reduced  hemoglobin.  Its  oxygen  is  in  more 
stable  combination,  however,  than  is  the  case  with  the  former  compound. 

Estimation  of  Hemoglobin. — The  most  exact  method  is  by  the  esti- 
mation of  the  amount  of  iron  (dry  hemoglobin  containing  0.42  per  cent, 
of  iron)  in  a  given  specimen  of  blood,  but  as  this  is  a  somewhat  complicated 
process,  various  methods  have  been  proposed  which,  though  not  so  exact, 
have  the  advantage  of  simplicity.  Of  the  several  varieties  of  hemo- 
globinometer,  one  of  the  oldest  adapted  to  its  purpose  is  that  invented  by 
professor  Fleischl,  of  Vienna.  In  this  instrument  the  amount  of  hemo- 
globin in  a  solution  of  blood  is  estimated  by  comparing  a  stratum  of 
diluted  blood  with  a  standard  solid  substance  of  uniform  tint  similar 
spectroscopically  to  diluted  blood.  The  instrument  has  been  modified 
and  made  more  accurate  by  Miescher.  The  Fleischl-Miescher  apparatus 
consists  of  a  stand  with  a  metal  plate  having  a  circular  opening  and  a 


ESTIMATION    OF    HEMOGLOBIN 


143 


plaster  mirror  below,  5,  figure  122,  which  casts  light  through  the  opening. 
Beneath  the  plate  is  a  metal  framework  containing  a  colored  glass  wedge, 
and  along  the  side  of  the  same  is  a  scale  graduated  so  as  to  indicate  the 
percentage  of  hemoglobin  corresponding  to  the  shades  of  the  different  parts 
of  the  wedge.  This  frame-  ^  ar 

work  can  be  moved  by  the 
wheel  T  which  fits  into  a  rack 
on  its  lower  surface.  The 
scale  can  be  read  through  a 
small  opening  M  in  the  plate. 
Into  the  large  circular  open- 
ing of  the  plate  fits  a  cylin- 
drical metal  cell  G  with  a  glass 
bottom  and  divided  by  a  metal 
partition  into  two  equal  parts. 
One  of  these  halves  lies  over 
the  wedge  and  is  filled  with 
distilled  water.  The  other 
contains  the  solution  of  blood 

in    which    the   hemoglobin  is 

j       ,-,,  FIG.  122. — Fleischl's  Hemoglobinometer. 

to  be  estimated.  The  appar- 
atus is  usually  supplied  with  three  cells.  Of  these,  the  first  two  are 
used  in  estimating  the  hemoglobin  according  to  Miescher.  This  is  the 
method  now  generally  used.  These  cells  are  furnished  with  a  glass  cover 
having  a  groove  whicn  fits  upon  the  partition  of  the  cell.  Over  this  cover 
is  placed  a  diaphragm  with  a  longitudinal  slit,  which  only  permits  of 
the  central  part  of  each  side  of  the  cell  being  seen. 

The  patient's  ear  or  finger  is  pricked,  and  the  blood  from  the  wound 
sucked  up  into  the  graduated  pipet  until  it  reaches  the  mark  J,  §,  or  -f,  a 
i  per  cent,  solution  of  sodium  carbonate  is  then  sucked  in  until  the  upper 
mark  is  reached.  The  pipet  is  then  well  shaken  in  order  to  mix  the  blood 
thoroughly.  One-half  of  each  of  the  two  cells,  which  are,  respectively,  12 
and  15  millimeters  high,  is  then  filled  with  the  mixture,  the  other  half 
being  filled  with  water.  An  important  point  is  that  the  liquids  should  com- 
pletely fill  the  cells.  The  cover-glasses  and  diaphragms  are  then  applied 
,  and  the  cells  are  ready  for  examination.  This  must  be  done  by  artificial 
light.  Moreover,  in  order  to  have  accurate  results,  light  of  the  same  inten- 
sity should  be  always  used.  One  of  the  cells  is  placed  on  the  plate  and  the 
wheel  T  turned  until  the  colors  of  the  two  halves  exactly  correspond.  When 
this  point  is  reached,  the  result  is  read  off  on  the  scale  through  the  opening 
M.  This  should  be  repeated  several  times  with  each  of  the  cells,  and  the 
average  of  the  readings  taken.  The  result  obtained  with  the  1 2-millimeter 


144  THE    BLOOD 

cell  should  be  multiplied  by  f  to  bring  it  up  to  that  of  the  larger.  For  example, 
suppose  the  result  of  several  readings  to  be: 

With  the  large  cell  (15  mm.) 54 .  oo 

With  the  small  cell  (12  mm.) 42  .00 

If  the  readings  obtained  with  the  large  cell  are  exactly  correct,  then  the  read- 
ings with  the  smaller  one  should  be  43.2,  since  54  X  £  =43.2.  Or,  if  the 
readings  with  the  smaller  cells  are  exact,  the  readings  with  the  larger  should 
be  52. 5, since 42  Xf  =52.5.  Hence  the  mean  of  54  and  52.5,  namely  53.25, 
should  be  taken  as  the  correct  figure.  On  looking  at  the  corrected  table  of 
hemoglobin  values  supplied  with  each  instrument,  we  would  find  that  this 
number  on  the  scale  corresponds  to  a  solution  containing  400  milligrams 
of  hemoglobin  per  1000  cubic  centimeters  of  solution.  But  our  original  dilu- 
tion was  either  i :  200,  i :  300,  or  i :  400,  according  as  our  pipet  had  been 
filled  with  blood  up  to  the  mark  -}-,  §,  or  J;  so  that  in  order  to  obtain  the  actual 
percentage  of  hemoglobin  in  the  blood  under  examination  we  should  be 
obliged  to  multiply  our  results  by  200,  300,  or  400.  In  the  example  we  have 
taken,  the  amount  of  hemoglobin  would  be,  if  our  dilution  was  i :  200, 400  X 
200=80,000  milligrams  =  80  grams  in  1,000  cubic  centimeters  =  8  grams 
in  100  cubic  centimeters,  or  8  per  cent. 

The  Dare's  hemoglobinometer  avoids  the  error  of  diluting  blood  by 
comparing  undiluted  blood  under  artificial  light  with  a  colored  scale  which 
is  graduated  after  standardization  against  a  hemoglobin  content  of  normal 
blood,  i.e.,  13.77  grams  of  hemoglobin  per  100  cubic  centimeters.  The 
instrument,  see  figure  1220,  consists  of  a  blood  pipette,  a  case  inclosing  the 
color  comparison  disc,  and  is  provided  with  a  small  telescope  for  reading 
the  color  contrasts  against  an  artificial  candle  light.  A  drop  of  blood  is 
drawn  with  a  lancet  from  a  finger  tip  or  the  lobe  of  the  ear,  is  allowed  to 
run  directly  between  the  plates  of  the  pipette  where  it  spreads  by  capil- 
larity. The  gradations  on  the  comparison  scale  are  read  under  candle 
light  and  the  computations  for  the  percentage  of  hemoglobin  in  the  sample 
made  against  the  normal.  Its  ease  of  manipulation  and  comparative 
accuracy  gives  to  this  method  the  status  of  a  clinical  favorite.  Another 
clinical  method  somewhat  less  readily  manipulated  is  that  of  Sahli. 

The  Talquist  method  enables  one  to  make  a  quick  approximation  of 
the  hemoglobin  content.  It  is  valuable  as  a  preliminary  test  to  the  Dare 
and  is  sufficiently  accurate  for  many  clinical  determinations. 

This  consists  of  a  series  of  shades  of  color  corresponding  to  undiluted 
blood  of  various  hemoglobin  values,  ranging  from  10  to  100  per  cent,  of  an 
arbitrary  scale.  This  scale  is  included  in  a  book,  the  remaining  pages  of 
which  consist  of  filter-paper,  which  is  used  for  absorbing  the  specimen  of 
blood  whose  hemoglobin  percentage  is  to  be  estimated.  The  blood- 
stained filter-paper  is  compared  with  the  hemoglobin  scale  by  direct  day- 


DERIVATIVES    OF    HEMOGLOBIN 


light  until  a  shade  is  found  with  which  it  corresponds.     For  approxi- 
mate results  this  method  has  proved  very  satisfactory. 


H 

FIG.  1220. — Dare  Hemoglobinometer.  Instrument  ready  for  use,  illustration  one- 
half  actual  size.  R,  Milled  wheel  by  which  the  color  prism  is  rotated  by  friction  exerted 
upon  its  edge.  S,  Case  enclosing  color  prism,  showing  stage  upon  which  the  blood 
pipet  slides.  T,  Movable  wing  pivoted  to  case.  When  drawn  outward  screens  the 
eyes  of  observer  from  the  light.  When  not  in  use  lies  superimposed  upon  the  circular 
prism  case,  occupying  no  extra  space.  U,  Telescoping  camera  tube  in  position  for 
examination.  V,  Opening  in  prism  case,  admitting  light  for  illumination  of  color  prism. 
The  white  glass  disc  of  prism  is  seen  inside.  W,  White  glass  of  blood  pipet.  X,  Pipet 
clamp  held  in  position  on  the  stage  by  grooves  and  guides.  F,  Detachable  candle- 
holder.  Z,  Rectangular  opening  in  edge  of  case  for  reading  hemoglobin  percentage 
indicated  by  beveled  blade.  L,  Light  (candle  or  electric.)  Color  prism.  E,  Prism  of 
colored  glass.  F,  Semi-circle  of  white  glass,  the  edge  carrying  the  index  of  hemoglobin 
percentage  in  black;  this  edge  also  serves  as  a  friction  surface  for  the  rubber-covered  roller 
by  which  the  prism  is  rotated.  G,  Hole  in  which  hub  is  fixed.  H,  Index  of  hemoglobin 
percentage  etched  in  black.  7,  Disc  of  white  glass  which  serves  to  break  the  glare  of 
direct  light  and  furnishes  a  white  background  to  view  the  shades  of  color. 

Derivatives  of  Hemoglobin. — Hematin. — By  the  action  of  heat  or 
of  acids  or  alkalies  in  the  presence  of  oxygen,  hemoglobin  can  be  split  up 
into  a  substance  called  hematin,  which  contains  all  the  iron  of  the  hemo- 
globin from  which  it  was  derived,  and  a  protein  residue,  a  histone,  globin. 
If  there  be  no  oxygen  present,  instead  of  hematin  a  body  called  hemochro- 
mogen  is  produced,  which,  however,  will  speedily  undergo  oxidation  into 
hematin. 

Hematin  is  a  dark  brownish  or  black  non-crystallizable  substance  of 
metallic  luster.  Its  percentage  composition  is  C,  64.30;  H,  5.50;  N,  9.06; 
Fe,  8.82;  O,  12.32;  which  gives  the  formula  C68H70N8Fe2O10  (Hoppe- 
Seyler).  It  is  insoluble  in  water,  alcohol,  and  ether;  soluble  in  the  caustic 
alkalies;  soluble  with  difficulty  in  hot  alcohol  to  which  is  added  sulphuric 
acid.  The  iron  may  be  removed  from  hematin  by  heating  it  with  fuming 


146  THE    BLOOD 

hydrochloric  acid  to  160°  C.,  and  a  new  body,  hematoporphyrin,  the  so-called 
iron-free  hematin,  is  produced.  Hematoporphyrin  (C88H74N8O12,  Hoppe- 
Seyler)  may  also  be  obtained  by  adding  blood  to  strong  sulphuric  acid,  and 
if  necessary  filtering  the  fluid  through  asbestos.  It  forms  a  fine  crimson 
solution,  which  has  a  distinct  spectrum,  viz.,  a  dark  band  just  beyond  D, 
and  a  second  all  but  midway  between  D  and  E.  It  may  be  precipitated  from 


FIG.  123.— Hematoidin  Crystals.     (Frey.)  FlG.  I23a.— Hemin  Crystals.     (Frey.) 

its  acid  solution  by  adding  water  or  by  neutralization,  and  when  redissolved 
in  alkalies  presents  four  bands,  a  pale  band  between  C  and  D,  a  second 
between  D  and  E,  nearer  D,  another  nearer  E,  and  a  fourth  occupying  the 
chief  part  of  the  space  between  b  and  F. 

Hematin  in  Acid  Solution. — If  an  excess  of  acetic  acid  is  added  to  blood, 
and  the  solution  boiled,  the  color  alters  to  brown  from  decomposition  of 
hemoglobin  and  the  setting  free  of  hematin;  by  shaking  this  solution  with 
ether,  a  solution  of  hematin  in  acid  solution  is  obtained.  The  spectrum  of 
the  ethereal  solution  shows  no  less  than  four  absorption  bands,  viz.,  one  in 
the  red  between  C  and  D,  one  faint  and  narrow  close  to  D,  and  then  two 
broader  bands,  one  between  D  and  E,  and  another  nearly  midway  between 
b  and  F.  The  first  band  is  by  far  the  most  distinct,  and  the  acid  aqueous 
solution  of  hematin  shows  it  plainly. 

Hematin  in  Alkaline  Solution. — If  a  caustic  alkali  is  added  to  blood  and 
the  solution  is  boiled,  alkaline  hematin  is  produced,  and  the  solution  becomes 
olive-green  in  color.  The  absorption  band  of  the  new  compound  is  in  the 
red,  near  to  D,  and  the  blue  end  of  the  spectrum  is  absorbed  to  a  considerable 
extent.  If  a  reducing  agent  be  added,  two  bands  resembling  those  of  oxy- 
hemoglobin,  but  nearer  to  the  blue,  appear;  this  is  the  spectrum  of  reduced 
hematin,  or  hemochromogen.  On  violently  shaking  the  reduced  hematin 
with  air  or  oxygen  the  two  bands  are  replaced  by  the  single  band  of  alkaline 
hematin. 

Hematoidin. — This  substance  is  found  in  the  form  of  yellowish  crystals, 
figure  123,  in  old  blood  extravasations  and  is  derived  from  the  hemoglobin. 
Their  crystalline  form  and  the  reaction  they  give  with  fuming  nitric  acid 
seem  to  show  them  to  be  closely  allied  to  bilirubin,  the  chief  coloring  matter 
of  the  bile,  and  in  composition  they  are  probably  either  identical  or  isomeric 
with  it. 

Hemin. — One  of  the  most  important  derivatives  of  hematin  is  hemin. 
It  is  usually  called  hydrochloride  of  hematin,  but  its  exact  chemical  com- 


DERIVATIVES  OF  HEMOGLOBIN  147 

position  is  uncertain.  Its  formula  is  said  to  be  C32H30N4FeO3HCl,  and  it 
contains  5.18  per  cent,  of  chlorine,  but  by  some  it  is  looked  upon  as  simply 
crystallized  hematin.  Although  difficult  to  obtain  in  bulk,  a  specimen  may 
be  easily  made  for  the  microscope  in  the  following  way:  A  small  drop  of 
dried  blood  is  finely  powdered  with  a  few  crystals  of  common  salt  on  a  glass 

Hemoglobin        +        Oxygen        «z±        Oxyhemoglobin  -*.      Methemoglobin 

on  heating  with  XV  on  heating  with        on  treating  with 

acidulated  alcohol  /       \  acidulated  alcohol       potassium  ferri- 

cyanide,  etc. 


Globin 


Hemochromogen  Hematin  Globin 

with  strong      /  \        heat  with 
sulphuric  and     /       \      glacial  acetic 
hydrobromic  acids   /  \    acid  and  sodium 

/  \chloride 

/  * 

Hematoporphyrin  Hemin 

(Iron-free  hematin.  (Hematin  hydrochloride) 

isomeric  or  identical 
with  bilirubin.) 

on  reduction,  with  Stoke's  reagent,  etc. 


Scheme  to  Show  the  Relations  of  Hemoglobin  and  its  Derivatives. 

slide  and  spread  out;  a  cover-glass  is  then  placed  upon  it,  and  glacial  acetic 
acid  added  by  means  of  a  capillary  pipet.  The  blood  at  once  turns  a  brown- 
ish color.  The  slide  is  then  heated,  and  the  acid  mixture  evaporated  to 
dryness  at  a  high  temperature.  The  excess  of  salt  is  washed  away  with 
water  from  the  dried  residue,  and  the  specimen  may  then  be  dried  and 
mounted.  A  large  number  of  small,  dark,  reddish-black  crystals  of  a  rhom- 
bic shape,  sometimes  arranged  in  bundles,  will  be  seen  if  the  slide  be  sub- 
jected to  microscopic  examination,  figure  123%. 

The  formation  of  these  hemin  crystals  is  of  great  interest  and  importance 
from  a  medico-legal  point  of  view,  as  it  constitutes  the  most  certain  and 
delicate  test  we  have  for  the  presence  of  blood  (not  of  necessity  the  blood 
of  man)  in  a  stain  on  clothes,  etc.  It  exceeds  in  delicacy  even  the  spectro- 
scopic  test.  Compounds  similar  in  composition  to  hemin,  but  containing 
hydrobromic  or  hydriodic  acid,  instead  of  hydrochloric,  may  be  also  readily 
obtained. 

Variations  in  the  Composition  of  Healthy  Blood. — The  conditions 
which  appear  most  to  influence  the  composition  of  the  blood  in  health  are 
these:  Diet,  Exercise,  Sex,  Pregnancy,  and  Age. 

Sex. — The  blood  of  men  differs  from  that  of  women,  chiefly  in  being  of 
somewhat  higher  specific  gravity,  from  its  containing  a  relatively  larger 
quantity  of  red  corpuscles. 

Pregnancy. — The  blood  of  pregnant  women  has  rather  lower  than  the 
average  specific  gravity.  The  quantity  of  the  colorless  corpuscles  is  in- 
creased in  the  later  months,  especially  in  primiparae;  it  is  also  claimed  that 
the  fibrin  is  increased  in  amount. 


148  THE    BLOOD 

Age. — The  blood  of  the  fetus  is  very  rich  in  solid  matter,  and  especially 
in  colored  corpuscles;  and  this  condition,  gradually  diminishing,  continues 
for  some  weeks  after  birth.  The  quantity  of  solid  matter  then  falls  during 
childhood  below  the  average,  rises  during  adult  life,  and  in  old  age  falls  again. 

Diet. — Such  differences  in  the  composition  of  the  blood  as  are  due  to  the 
temporary  presence  of  various  matters  absorbed  with  the  food  and  drink, 
as  well  as  the  more  lasting  changes  which  must  result  from  generous  or  poor 
diet,  respectively,  need  be  here  only  referred  to. 

Effects  of  Bleeding. — The  result  of  bleeding  is  to  diminish  the  specific 
gravity  of  the  blood,  and  so  quickly  that  in  a  single  venesection  the  portion 
of  blood  last  drawn  has  often  a  less  specific  gravity  than  that  of  the  blood  that 
flowed  first.  This  is,  of  course,  due  to  absorption  of  fluid  from  the  tissues 
of  the  body.  The  physiological  import  of  this  fact,  namely,  the  instant 
absorption  of  liquid  from  the  tissues,  is  the  same  as  that  of  the  intense  thirst 
which  is  so  common  after  either  loss  of  blood  or  the  abstraction  from  it  of 
watery  fluid,  as  in  cholera,  diabetes,  and  the  like. 

For  some  little  time  after  bleeding,  the  want  of  colored  corpuscles  is  well 
marked,  but  with  this  exception  no  considerable  alteration  seems  to  be 
produced  in  the  composition  of  the  blood  for  more  than  a  very  short  time; 
the  loss  of  the  other  constituents,  including  the  colorless  corpuscles,  being 
very  quickly  repaired. 

Variations  in  Different  Parts  of  the  Body. — The  composition  of  the  blood, 
as  might  be  expected,  is  found  to  vary  in  different  parts  of  the  body.  Thus 
arterial  blood  differs  from  venous;  and  although  its  composition  and  general 
characters  are  uniform  throughout  the  whole  course  of  the  systemic  arteries, 
they  are  not  so  throughout  the  venous  system,  the  blood  contained  in  some 
veins  differing  markedly  from  that  in  others. 

Differences  between  Arterial  and  Venous  Blood. — The  differences  between 
arterial  and  venous  blood  are  these: 

Arterial  blood  is  bright  red,  from  the  fact  that  almost  all  its  hemoglobin 
is  combined  with  oxygen,  oxyhemoglobin,  while  the  dark  red  tint  of  venous 
blood  is  due  to  the  deoxidation  of  a  certain  quantity  of  its  oxyhemoglobin, 
and  its  consequent  reduction  to  the  hemoglobin. 

Arterial  blood  coagulates  somewhat  more  quickly. 

Arterial  blood  contains  more  oxygen  than  venous  and  less  carbon 
dioxide  gas. 

Some  of  the  veins  contain  blood  which  differs  from  the  ordinary  standard 
considerably.  These  are  the  portal,  the  hepatic,  and  the  splenic  veins. 

Portal  Blood. — The  blood  which  the  portal  vein  conveys  to  the  liver  is 
supplied  from  two  chief  sources;  namely,  from  the  gastric  and  mesenteric 
veins,  which  contain  the  soluble  elements  of  food  absorbed  from  the  stomach 
and  intestines  during  digestion,  and  from  the  splenic  vein.  It  must,  there- 
fore, combine  the  qualities  of  the  blood  from  each  of  these  sources. 


GLOBULOCIDAL    AND    OTHER    PROPERTIES    OF    SERUM  149 

The  blood  in  the  gastric  and  mesenteric  veins  will  vary  much  according 
to  the  stage  of  digestion  and  the  nature  of  the  food  taken,  and  can  therefore 
be  seldom  exactly  the  same.  Speaking  generally  and  without  considering 
the  sugar  and  other  soluble  matters  which  may  have  been  absorbed  from 
the  alimentary  canal,  this  blood  appears  to  be  deficient  in  solid  matters, 
especially  in  colored  corpuscles,  owing  to  dilution  by  the  quantity  of  water 
absorbed,  to  contain  an  excess  of  protein  matter,  and  to  yield  a  less  tenacious 
kind  of  fibrin  than  that  of  blood  generally. 

The  blood  of  the  portal  vein,  combining  the  peculiarities  of  its  two  factors 
the  splenic  and  mesenteric  venous  blood,  is  usually  of  lower  specific  gravity 
than  blood  generally,  is  more  watery,  contains  fewer  colored  corpuscles, 
more  proteins,  and  yields  a  less  firm  clot  than  that  yielded  by  other  blood, 
owing  to  the  deficient  tenacity  of  its  fibrin. 

Guarding  (by  ligature  of  the  portal  vein)  against  the  possibility  of  an 
error  in  the  analysis  from  regurgitation  of  hepatic  blood  into  the  portal  vein, 
recent  observers  have  determined  that  hepatic  venous  blood  contains  less 
water,  proteins,  and  salts  than  the  blood  of  the  portal  veins;  but  that  it 
yields  a  much  larger  amount  of  extractive  matter,  in  which  is  one  constant 
element,  namely,  glucose,  which  is  found  whether  carbohydrates  have 
been  present  in  the  food  or  not. 

GLOBULOCIDAL    AND    OTHER    PROPERTIES    OF    SERUM. 


Cytolysis. — It  has  been  known  for  some  time  that  the  sera  of  certain 
animals  when  injected  into  the  circulation  of  animals  of  another  species  will 
cause  destructive  changes  in  the  blood  corpuscles,  accompanied  by 
symptoms  of  poisoning,  which  may  even  end  fatally.  These  results  served 
to  bring  into  disrepute  the  use  of  foreign  blood  in  transfusion,  which  has 
consequently  been  practically  abandoned.  The  discharge  of  the  hemo- 
globin of  the  red  blood  corpuscles  and  their  solution  in  the  plasma  (laking) 
is  now  included  in  the  general  term  Cytolysis,  and  more  specifically  known 
as  Hemolysis.  Agents  which  produce  such  an  effect  are  known  as  hemolytic 
or  hemotoxic  agents. 

Transfusion  of  the  blood  of  one  animal  into  the  vessels  of  another  is 
often  quickly  fatal  because  of  the  hemolytic  reactions  of  the  two  bloods. 
Transfusions  between  different  species  or  distantly  related  animals  are  as  a 
rule  not  possible.  But  the  blood  of  different  individuals  of  the  same 
species  are  usually  not  lytic.  This  subject  possesses  great  importance 
to  man  because  of  the  growing  practice  of  blood  transfusions  in  man. 
However  not  all  human  bloods  can  be  blended  without  great  danger  though 
as  a  rule  members  of  the  same  family  are  miscible.  Human  bloods  must 
first  be  tested  or  typed,  as  it  is  called,  to  determine  whether  or  not  lysis 


150  THE    BLOOD 

would  occur  on  transfusion.  Four  human  types  have  been  described  by 
serologists.  No  transfusion  is  now  ever  performed  without  this  pre- 
liminary test  which  is  absolutely  necessary  lest  immediate  solution  of  the 
red  corpuscles  be  produced  and  the  death  of  the  recipient  follow. 

The  serum  of  one  animal  may  be  made  to  acquire  lytic  properties  for 
the  blood  of  another.  This  adaptation  is  brought  about  in  the  following 
way:  For  instance,  the  blood  of  the  guinea-pig,  which  is  not  normally 
lytic  for  the  red  cells  of  the  rabbit,  may  be  adapted  to  the  latter  by  pre- 
viously, at  several  successive  intervals  (three  to  seven  days)  injecting  into 
the  abdominal  cavity  or  subcutaneous  tissues  of  the  guinea-pig  small 
quantities  of  rabbit's  blood.  If  now  a  small  quantity  of  serum  be  obtained 
from  the  guinea-pig  by  the  usual  methods  and  mixed  in  a  test-tube  with 
some  of  the  rabbit's  blood  diluted  with  physiological  salt  solution,  hemoly- 
sis  occurs;  that  is,  the  coloring  matter  of  the  rabbit's  red  blood-cells  goes 
into  solution  and  the  cells  appear  under  the  microscope  as  shadow  cor- 
puscles or  ghosts,  devoid  of  hemoglobin.  Such  an  artificially  produced 
hemolytic  serum  is  only  lytic  for  the  blood  of  the  animal  species  for  which 
it  has  been  adapted.  It  is  true  that  it  may  also  show  slightly  lytic 
properties  for  closely  allied  species.  It  has  therefore  been  suggested  as  a 
possible  valuable  aid  in  determining  relationships  of  various  animal 
species. 

Concerning  the  nature  of  the  lytic  substance,  it  has  been  found  that  it 
probably  consists  of  two  bodies  acting  conjointly,  for  if  the  serum  be  heated 
to  56°  C.  for  a  short  time,  its  lytic  powers  are  lost,  but  may  be  restored  by 
adding  a  little  serum  of  another  animal  of  the  same  species  which  has  not 
been  adapted,  and  whose  serum  is  consequently  not  in  itself  lytic.  Of  these 
two  bodies,  therefore,  one  is  stable  and  is  formed  only  in  the  adapted  serum, 
while  the  other  is  more  unstable  or  labile,  destroyed  at  56°  C.,  and  exists 
normally  in  the  blood  plasma.  The  former  is  known  as  the  immune  body, 
the  amboceptor,  and  the  latter  as  the  alexin,  or  complement.  Lysis  occurs 
only  when  both  are  present  at  the  same  time,  and  not  through  the  agency  of 
one  or  the  other  singly. 

This  cytolytic  adaptation  has  been  extended  to  include  other  cells  besides 
the  red  blood  corpuscles.  Thus  in  a  similar  manner  leucolytic,  hepatolytic, 
nephrolytic,  and  a  number  of  other  lytic  sera  have  been  developed. 

It  is  further  possible,  under  certain  circumstances,  that  substances  may 
be  developed  in  the  tissues  which  are  lytic  for  other  tissue  cells  of  the  same 
animal,  autolytic  substances.  This  may  be  a  physiologically  important  process 
in  the  elimination  of  worn-out  tissue  cells,  cellular  elements  in  injury,  in- 
flammation, etc. 

Agglutinative  Substances. — A  further  property  of  adapted  sera  is 
that  of  agglutination.  The  adaptation  is  secured  in  the  same  way  as  in 


GLOBULOCIDAL  AND  OTHER  PROPERTIES  OF  SERUM     151 

the  production  of  cytolysins.  In  fact,  both  cytolysis  and  agglutination  may 
occur  at  the  same  time.  The  normal  blood  serum  of  some  animals  may 
be  agglutinative  for  the  blood-cells  of  some  other  species.  In  normal  serum, 
agglutinative  and  cytolytic  properties  may  be  present  together  or  one  only 
may  be  normally  present. 

The  activity  of  agglutinative  substances  is  not  destroyed  at  a  tempera- 
ture of  56°  C.  They  do  become  inert,  however,  at  70°  C.,  and,  furthermore, 
they  cannot  be  restored  by  adding  normal  serum,  as  is  the  case  with  cytolysins. 

Hemagglutinative  substances  are  found  in  certain  plant  seeds;  e.g.,  in 
castor  oil  beans  (Ricinus  communis) ,  in  cotton  seed,  and  in  the  legumes. 

Precipitins. — Other  forms  of  adaptive  substances  which  may  be  found 
in  animal  serum  are  those  which,  when  mixed  with  the  substances  by  means 
of  which  adaptation  has  been  secured,  form  a  precipitate.  By  this  means 
blood  of  different  species  of  animals  may  be  detected  even  when  in  a  dried 
state.  It  has  been  suggested  as  a  possible  valuable  aid  in  medico-legal 
cases,  since  human  blood  in  a  dilution  of  i  to  50,000  has  been  recognized 
by  this  means. 

Opsonins. — Wright  and  Douglass  have  shown  that  there  are  certain 
substances  in  the  serum  that  affect  bacteria  in  such  a  way  that  they  are  more 
easily  taken  up  and  destroyed  by  leucocytes.  The  phagocytic  power  of  the 
leucocytes  in  destroying  toxic  bacteria  is  not  made  to  increase  by  stimulative 
substances,  as  Metchnikoff  believed,  but  rather  by  those  materials  in  the 
serum  diminishing  the  resisting  power  of  the  bacteria.  These  substances 
are  called  by  their  discoverers  opsonins.  They  found  opsonin  present  in 
normal  serum,  but  also  found  that  its  quantity  varies  under  certain  conditions. 
They  suggested  that  the  opsonins  could  be  measured  by  determining  the 
phagocytic  power.  The  ratio  of  the  average  number  of  bacteria  taken  up  by 
leucocytes  in  normal  serum  to  the  number  taken  up  in  the  immune  serum, 
they  called  the  opsonic  index. 

Antitoxins. — Certain  kinds  of  bacteria,  notably  the  diphtheria  and 
tetanus  organisms,  elaborate  poisonous  substances  known  as  toxins.  The 
pathological  conditions  resulting  from  such  infections  are  produced  by  the 
poisons  so  formed.  Behring  first  showed  that  immunity  to  diphtheria  was 
due  to  the  presence  in  the  blood  plasma  and  blood  serum  of  substances  which 
apparently  combine  with  and  so  prevent  the  toxic  action  of  the  bacterial 
products.  This  antitoxic  power  of  the  blood  can  be  artificially  developed  by 
injecting  small  doses  of  the  toxins  into  an  animal,  usually  a  horse,  at  inter- 
vals of  some  days.  The  protective  power  of  the  blood  against  the  toxins  can 
thus  be  developed  to  a  relatively  enormous  degree.  The  serum  of  an  arti- 
ficially immunized  animal  can  be  injected  into  other  individuals  of  the  same 
or  other  species  and  an  immunity  will  be  conferred  on  the  person  or  animal 
so  treated.  Similarly,  the  antitoxic  sera  have  a  curative  effect  in  infected 
individuals  if  the  disease  is  not  too  far  advanced.  Antitoxic  sera  are  specific 


THE    BLOOD 

for  the  particular  toxin  used  for  the  immunization.  Antitoxins  can  be 
similarly  prepared  for  the  naturally  occurring  vegetable  toxins,  ricin  and 
abrin,  for  snake  venoms,  etc. 

Anaphylaxis. — It  has  been  found  that  if  an  animal,  especially  the 
guinea  pig,  be  injected  with  even  the  minutest  quantity  of  protein  material, 
that  after  ten  to  fourteen  days,  the  animal  becomes  susceptible  to  a  sec- 
ond injection  of  the  same  material  if  made  intravascularly.  Thus  a  guinea 
pig  may  be  sensitized  with  o.oooooi  of  a  c.c.  of  blood  serum;  after  two 
weeks  have  elapsed,  the  introduction  of  a  half  c.c.  of  the  same  serum  into 
the  circulation  will  in  a  few  minutes  lead  to  respiratory  failure  and  death. 
This  phenomenon  is  relatively  specific  for  foreign  protein  substances. 
Guinea-pigs  which  have  recovered  from  the  second  injection  acquire  a 
temporary  immunity  against  a  third  injection  of  the  same  protein.  It  has 
been  found  that  this  peculiar  susceptibility  is  transferred  from  the  mother 
guinea-pig  to  successive  litters. 

Nature  of  the  Antisubstances  in  Blood. — The  lecithins  and  fatty  acids, 
especially  oleic  acid,  will  in  a  measure  replace  a  hemolytic  complement. 
The  antitoxins  and  agglutinins  in  the  blood  seem  either  to  be  associated 
with,  or  actually  are  a  portion  of,  the  para-  or  pseudoglobulin.  During  im- 
munization, the  pseudoglobulin  of  the  blood  may  be  twice  the  normal  con- 
tent; coincidently  the  per  cent,  of  the  serumalbumin  will  diminish.  The 
protein  changes  in  the  blood  of  horses  and  the  antitoxic  variations,  how- 
ever, are  not  parallel,  and  no  quantitative  relationship  has  been  established. 

PHYSICAL  FACTORS  OF  BLOOD  PLASMA  OR  SERUM. 

Diffusion,  Osmosis,  Dialysis. — The  term  diffusion  has  long  been  ap- 
plied to  the  regular  mixing  of  the  molecules  of  two  gases  when  brought 
into  contact  in  a  confined  space,  this  interpenetration  being  due  to  the 
to-and-fro  movements  of  their  molecules.  More  recently  it  has  been 
applied  to  the  mixing  of  the  molecules  of  two  solutions  when  brought  into 
contact,  as  it  has  been  found  that  they  act  in  the  same  way  and  obey  the 
same  laws  as  gases.  If,  however,  the  two  solutions  are  separated  by  a 
membrane,  permeable  to  the  solutions,  diffusion  will  still  occur.  To  this 
form  of  diffusion  the  term  osmosis  has  been  applied  in  the  case  of  water, 
and  dialysis  in  the  case  of  diffusible  substances.  All  bodies  can  be 
divided  into  two  groups,  crystalloids  and  colloids.  To  the  former  group 
belong  bodies  having  a  crystalline  form,  which  readily  go  into  solution 
in  water.  All  such  bodies  are  diffusible  (dialyzable),  their  power  of 
dialysis,  however,  varying  considerably.  To  the  second  group  belong 
such  bodies  as  have  no  crystalline  form  (amorphous).  These  are  generally 
bodies  with  a  large  molecule,  which  form  colloidal  suspensions  in  water  and 
are  only  slightly  or  not  at  all  diffusible.  An  exception  to  the  first  group 
is  hemoglobin,  which  has  a  very  large  molecule,  and  is  crystalline  but  is 


PHYSICAL  FACTORS  OF  BLOOD  PLASMA  OR  SERUM 


not  diffusible.     The  following  may  serve  as  simple  illustrations: 

Take  a  jar  and  divide  it  in  two  equal  parts  by  an  animal  membrane,  M, 
figure  124,  and  place  an  equal  amount  of  distilled  water  in  the  two  sides,  A 
and  B.  Now,  since  the  molecules  of  water  act  like  those  of  a  gas  and  are 
continually  moving  to  and  fro,  bombarding  all  the  surfaces  of  their  retainer, 
the  molecules  of  water  in  A  and  B  will  be  continually  striking  all  the  surfaces 
of  A  and  B;  but  since  the  membrane  is  permeable  to 
the  water  molecules,  there  will  be  a  continual  inter- 
change of  molecules  between  A  and  B.  If  now,  in  one 
side  A  we  place  a  solution  of  sodium  chloride,  still 
keeping  water  in  B,  the  membrane  being  permeable  to 
the  sodium  chloride,  the  first  thing  we  should  notice 
would  be  an  increase  in  the  amount  of  water  in  A.  For- 


M 


M 


— T 


FIG.  124.  FIG.  125. 

merly  it  would  have  been  said  that  "  the  salt  had  attracted  the  water."  Now 
we  should  say  that  the  salt  had  a  certain  osmotic  pressure.  The  salt,  how- 
ever, being  able  to  pass  (dialyse)  through  the  membrane,  will  do  so,  and  this 
will  continue  until  the  strength  of  the  two  salt  solutions,  and  therefore  the 
osmotic  pressure  on  both  sides,  is  equal. 

Osmotic  Pressure. — If  now  in  A  we  place  a  solution  of  some  soluble 
colloidal  substance  to  which  the  membrane  is  impermeable,  or  else  replace 
the  membrane,  M,  we  used  in  our  former  experiment  by  one  which  is  not 
permeable  to  the  sodium  chloride,  and  arrange  our  jar  as  in  figure  125,  so 
as  to  be  able  to  read  off  any  increase  of  water  which  may  pass  into  A,  we 
will  notice  that  the  amount  of  liquid  in  A  will  continue  to  increase  up  to  a 
certain  point.  Once  that  point  is  reached,  there  will  be  no  further  change, 
since  the  substance  in  solution,  in  A ,  cannot  pass  through  the  membrane  as 
in  the  previous  example.  This  pressure  can  be  measured  and  expressed  in 
millimeters  of  mercury.  It  is  constant  for  all  solutions  of  this  substance 
that  are  of  the  same  concentration  when  measured  under  like  conditions  of 
temperature  and  pressure,  and  is  called  the  osmotic  pressure  of  this  solution. 

Of  the  numerous  explanations  regarding  the  nature  of  osmotic  pressure 
which  has  been  more  or  less  satisfactory,  a  simple  one,  and  one  that  can 


154  THE    BLOOD 

be  easily  understood,  is  as  follows:  In  figure  125  one  surface  of  the  mem- 
brane is  being  bombarded  by  the  molecules  of  a  non-diffusible  substance 
mixed  with  those  of  a  diffusible  one  (water) ;  while  the  other  surface  is  being 
bombarded  entirely  by  water  molecules.  The  former  condition  permits 
only  a  fraction  of  the  molecules  to  diffuse  out,  since  fewer  water  molecules 
get  to  the  surface  of  the  membrane;  while  the  latter  permits  all  of  the 
molecules  which  reach  it  to  pass  through. 

Osmotic  pressure  can  be  estimated  in  several  different  ways  in  addition 
to  the  above,  viz.,  the  determination  of  the  freezing-point  of  the  solution, 
determination  of  the  boiling-point,  determination  of  the  electrical  conduc- 
tivity. The  results  obtained  with  the  various  methods  agree  very  closely. 
The  following  solutions  have  the  same  osmotic  pressure:  Sodium  chloride, 
i .  64  per  cent. ;  potassium  nitrate,  i .  09  per  cent. ;  sugar  5 . 5  per  cent. 

Isotonic  Solutions. — Solutions  that  have  the  same  osmotic  pressure 
are  called  isotonic.  The  term  isotonic  is  a  relative  one,  implying  a  compari- 
son with  some  other  solution  taken  as  a  standard.  In  physiology  it  has  been 
customary  to  take  blood  plasma  as  a  standard.  A  solution  of  o .  64  per  cent, 
sodium  chloride  is  isotonic  for  the  blood  plasma  of  the  frog,  and  a  o .  9  per 
cent,  solution  for  that  of  man.  Further,  any  solution  which  is  of  a  lower 
osmotic  pressure  than  the  standard  solution  is  said  to  be  hypoisotonic  (hypo- 
tonic)  in  relation  to  that  solution.  A  solution  of  a  higher  osmotic  pressure 
is  said  to  be  hyperisotonic  (hypertonic}. 

Water  passes  in  the  Direction  of  the  Arrows. 
Hypertonic  saline  solution (2  per  cent.) 

I 

Blood-plasma 

Ti 

Isotonic  saline  solution (o .  64  per  cent.) 

I. 

Hypotonic  saline  solution (o-3     per  cent.) 

If  a  hypotonic  solution  be  mixed  with  blood,  water  from  the  hypotonic 
•solution  passes  through  the  cell  membrane  of  the  red  corpuscles  into  the 
stroma  and  causes  it  to  swell.  The  hemoglobin  at  the  same  time  passes 
out  and  goes  into  solution  in  the  diluted  plasma.  On  the  other  hand,  the 
addition  of  a  hypertonic  solution  to  the  plasma  causes  the  red  corpuscles 
to  lose  their  water  and  become  crenated.  The  principles  of  osmosis  have 
been  derived  from  the  action  of  substances  separated  by  dead  animal  or 
plant  membranes.  It  must  be,  however,  remembered  that  in  the  applica- 
tion of  these  principles  to  processes  occurring  in  the  living  organism,  the 
cells,  forming  the  various  membanes,  are  an  important  modifying  factor. 
It  is  probable  that  physico-chemical  processes,  occurring  in  the  protoplasm 
of  the  cell,  may  change  its  permeability  to  the  same  substance  at  different 
times. 

THE  CHARACTER  AND  COMPOSITION  OF  LYMPH. 

The  lymph  is  the  fluid  which  immediately  surrounds  the  tissue  cells  of 


ANALYSIS    OF    LYMPH  155 

the  living  body.  It  fills  up  the  spaces  between  the  cells  themselves  and 
between  the  cells  and  the  blood  vessels  which  ramify  among  the  cell  masses. 
The  lymph,  therefore,  is  an  intermediate  fluid  between  blood  plasma  on  the 
one  hand  and  the  tissue  cells  on  the  other,  receiving  its  ingredients  by  the 
passage  of  fluid  from  the  plasma  through  the  walls  of  the  finer  blood  vessels 
in  the  one  direction  and  by  the  discharge  of  the  substances  from  the  cells 
themselves  in  the  other. 

The  Chemical  Composition  of  the  Lymph. — Since  the  chief  source 
of  the  lymph  is  the  blood  plasma,  one  would  naturally  expect  that  its  chemi- 
cal composition  would  be  very  similar  to  that  of  plasma,  which  is  in  fact  the 
case.  The  variations  that  are  noted  in  lymph  taken  from  definite  sources  no 
doubt  have  their  origin  in  the  fact  that  the  lymph  passes  through  these 
organs  slowly,  and  that  ingredients  peculiar  to  the  necessities  of  the  func- 
tion and  growth  of  the  differentiated  tissue  of  the  organ  are  taken  from  the 
lymph  in  special  organs.  Lymph  obtained  from  a  human  lymphatic  fistula 
has  been  analyzed;  the  figures  from  Hammarsten  are  as  follows,  though 
considerable  variations  appear  in  the  analyses  from  other  authorities: 

ANALYSIS  OF  LYMPH. 

Per  cent. 

Water 94-5      to  96 .  5 

Solids 3.5      to     5.5 

Proteins 3.4      to    4.1 

Ethereal  extract o .  06    to    0.13 

Sugar o.i 

Salts 0.8      to    0.9 

Sodium  chloride °-55    to    0.58 

Sodium  carbonate 0.24 

Disodic  phosphate 0.028 

The  most  notable  fact  to  be  derived  from  this  composition  table  is  the 
low  percentage  of  proteins  present  in  the  lymph. 

The  Formation  of  Lymph. — The  manner  in  which  the  substances  in 
the  lymph  pass  through  the  walls  of  the  capillaries  from  the  plasma  is  a 
question  which  has  been  surrounded  with  considerable  difficulty.  It  was 
thought  by  Ludwig  and  many  of  his  followers  that  the  process  involved 
is  merely  one  of  filtration.  Certainly  the  blood-pressure  in  the  capillaries 
is  in  the  main  greater  than  that  of  the  pressure  of  the  lymph  in  the  surround- 
ing tissues,  and  this  positive  pressure  will  contribute  so  much  to  the  direct 
ingredients  of  the  blood  plasma  through  the  capillary  walls.  It  is  true,  as  a 
matter  of  experiment,  that  anything  which  contributes  to  an  increase  in 
the  capillary  pressure  is  very  apt  to  produce  an  edema  of  the  corresponding 
tissues.  Since  the  colloidal  materials  represented  by  the  protein  are  non- 
diffusible,  one  would  by  this  theory  expect  to  find  a  diminished  percentage 
in  the  lymph,  which  is  true,  though  not  to  the  extent  which  the  theory 
demands. 


156  THE    BLOOD 

Heidenhain  was  the  first  to  question  the  adequacy  of  the  blood-pres- 
sure and  filtration  hypothesis.  He  showed  that  many  of  the  conditions 
under  which  lymph  formation  takes  place  are  not  sufficient  to  produce  filtra- 
tions  of  the  material  found.  He  advanced  the  hypothesis  that  the  living 
endothelial  lining  of  the  blood  vessels  exerted  a  secretory  activity  in  lymph 
production.  He  discovered  that  various  substances  known  as  lymphagogues 
when  introduced  into  the  circulatory  system  produce  a  remarkable  increase 
in  the  flow  of  lymph  from  the  thoracic  duct.  Further,  he  noticed  that  the 
concentration  of  the  lymph  was  changed;  i.e.,  increased.  Heidenhain 
thought  that  the  lymphagogues  acted  directly  on  the  capillary  and  endothelial 
lining  stimulating  these  cells  to  produce  a  greater  quantity  of  lymph.  He 
divided  such  substances  into  two  classes.  The  best  known  representatives 
of  the  " first  class"  are  such  as  proteoses  and  peptones,  leech  extract,  ex- 
tract of  crustacean  tissue,  etc.  The  lymphagogues  of  the  second  class  are 
the  neutral  inorganic  salts,  sugars,  and  other  crystalline  substances.  These 
all  cause  a  marked  flow  of  lymph.  The  lymphagogues  as  a  class  cause  fall 
of  blood-pressure;  for  example,  proteose-peptone  injections.  This  fact 
argues  against  their  purely  physical  action.  Many  drugs  act  to  increase 
the  flow  of  lymph  in  a  way  which  cannot  be  presumed  to  be  other  than  nor- 
mal; i.e.,  they  stimulate  the  physiological  processes  going  on  in  the  endo- 
thelial cells.  Such  observations  contribute  strongly  to  the  view  advanced 
by  Heidenhain.  Many  investigations  have  been  brought  to  the  support  of 
the  hypothesis  that  lymph  formation  is  largely  a  process  of  secretion,  yet 
it  seems  at  the  present  time  that  we  cannot  wholly  deny  that  filtration  and 
osmosis  play  a  part  in  the  processes. 

In  following  the  action  of  peptones  and  proteoses,  Pick  and  Spiro  came 
to  the  conclusion  that  it  was  not  peptone,  but  some  contaminating  substance 
which  produced  the  characteristic  action.  This  hypothetical  substance 
they  called  peptozyme.  Underhill  re-examined  the  influence  of  peptones, 
using  preparations  made  from  plant  proteins  by  hydration  with  enzymes, 
heat,  and  acid,  carrying  the  hydrolysis  to  a  greater  extent  than  did  Pick  and 
Spiro.  Underhill  still  obtained  the  great  increase  in  the  flow  of  lymph 
together  with  the  usual  fall  of  blood-pressure.  Mendel  published  the 
result  of  a  demonstration  of  post-mortem  lymph  flow  in  which  he  showed 
that ' '  the  lymph  continued  to  flow  for  four  hours  without  any  extraordinary 
mechanical  assistance"  after  the  death  of  the  animal.  This  observation 
would  seem  to  give  complete  refutation  of  the  filtration  hypothesis.  A 
similar  post-mortem  salivary  secretion  has  been  observed,  and  in  each  case 
the  processes  involved  must  be  assumed  to  be  physiological  rather  than 
purely  physical  phenomena.  Certainly  the  permeability  or  activity  of  the 
endothelial  lining  of  the  blood  vessels  varies  greatly  at  different  times  in 
the  life  of  an  individual,  and  this  variation  in  function  is  associated  with  the 
marked  change  in  the  character  and  quantity  of  lymph  produced. 


THE  RED  CORPUSCLES  157 

LABORATORY  EXPERIMENTS  FOR  THE  EXAMINATION  OF 

THE  BLOOD. 

1.  Microscopical  Examination  of  the  Blood. — Mount  a  drop  of  frog's 
blood  in  o .  7  per  cent,  sodium  chloride  and  examine  with  the  high  power  of  a 
compound  microscope.     The  red  corpuscles  will  appear  as  oval  nucleated 
discs  with  a  faint  yellowish  color,  figure  no.     Here  and  there  white  granular 
cells  of  irregular  outline  will  be  noted,  the  white  corpuscles.     Examine  the 
drop   of  blood   with   a  high  magnifying  power  (oil-immersion  lens)  and 
sketch  the  outline  of  the  blood-cells.     Select  the  white  corpuscle  which  is 
most  irregular  in  outline  and  make  a  series  of  outline  drawings  once  every 
minute  to  show  its  ameboid  movements,  figure  117. 

Draw  a  drop  of  your  own  blood  by  puncturing  the  tip  of  the  finger,  under 
sterile  conditions,  and  mount  in  a  drop  of  o .  9  per  cent,  physiological  saline. 
Examine  with  a  high  power,  note  the  small  biconcave  red  corpuscles  which 
appear  faintly  yellow  in  color  and  even  adhere  in  rouleaux,  figure  109.  The 
white  corpuscles  will  appear  as  somewhat  larger  granular  discs  differing  in 
form  and  size.  By  mounting  a  drop  of  blood  on  a  warm  stage  the  ameboid 
movements  of  the  white  corpuscles  can  be  observed  with  comparative  ease. 

2.  Action  of  Fluids  on  the  Red  Corpuscles. — Water. — When  water  is 
added  gradually  to  frog's  blood,  the  oval  disc-shaped  corpuscles  become 
spherical  and  gradually  discharge  their  hemoglobin,  a  pale,  transparent 
stroma  being  left  behind.     Human  red  blood-cells  change  from  a  discoidal 


FIG.  126.  FIG.  127.  FIG.  128.  FIG.  129. 

FIG.  126. — Effect  of  Hypertonic  Salt  Solution  on  the  Red  Blood  Corpuscles  of  Man. 
PIG.  127. — Effect  of  Acetic  Acid.  FIG.  128. — Effect  of  Tannin.  FIG.  129. — Effect 
of  Boric  Acid. 

to  a  spheroidal  form  and  discharge  their  cell  contents,  becoming  quite  trans- 
parent and  all  but  invisible  (ghost  corpuscles). 

Hypertonic  Salt  Solutions. — Mount  a  drop  of  human  blood  in  2  per  cent, 
sodium- chloride  solution.  The  red  blood-cells  lose  their  disc  shape  and  be- 
come spherical  with  spinous  projections  or  crenations,  figure  126. 

The  original  form  of  the  red  blood-cells  can  be  restored  by  transferring 
them  to  isotonic  salt  solution. 

Dilute  Acetic  Acid. — This  reagent  causes  the  nucleus  of  the  red  blood- 
cells  in  the  frog  to  become  more  clearly  defined;  if  the  action  is  prolonged, 
the  nucleus  becomes  strongly  granulated,  and  all  the  coloring  matter  seems 
to  be  concentrated  in  it,  the  surrounding  cell  substance  and  outline  of  the  cell 
becoming  almost  invisible;  after  a  time  the  cells  lose  their  color  altogether. 


158  THE    BLOOD 

The  cells  in  figure  127  represent  the  successive  stages  of  the  change.  A 
similar  loss  of  color  occurs  in  the  red  cells  of  human  blood,  which,  from  the 
absence  of  nuclei,  seem  to  disappear  entirely. 

Alkalies. — Alkalies  cause  the  red  blood  corpuscles  to  absorb  water  and 
finally  to  disintegrate. 

Chloroform  and  Ether. — These  reagents  when  added  to  the  red  blood- 
cells  of  the  frog  cause  them  to  part  with  their  hemoglobin;  the  stroma  of  the 
cells  becomes  gradually  broken  up.  A  similar  effect  is  produced  on  the 
human  red  blood-cell. 

Tannin  and  Boric  Acid. — When  a  2  per  cent,  fresh  solution  of  tannic  acid 
is  applied  to  frog's  blood  it  causes  the  appearance  of  a  sharply  defined  little 
knob,  projecting  from  the  free  surface  (Roberts'  macula}.  The  coloring 
matter  becomes  at  the  same  time  concentrated  in  the  nucleus,  which  grows 
more  distinct,  figure  128.  A  somewhat  similar  effect  is  produced  on  the 
human  red  blood  corpuscle. 

A  2  per  cent,  solution  of  boric  acid  applied  to  nucleated  red  blood-cells  of 
the  frog  will  cause  the  concentration  of  all  the  coloring  matter  in  the  nucleus; 
the  colored  body  thus  formed  gradually  quits  its  central  position,  and  comes 
to  be  partly,  sometimes  entirely,  protruded  from  the  surface  of  the  now 
colorless  cell,  figure  129.  The  result  of  this  experiment  led  Briicke  to  dis- 
tinguish the  colored  contents  of  the  cell  (zoo'id)  from  its  colorless  stroma 
(ecoid).  When  applied  to  the  non- nucleated  mammalian  corpuscle  its  effect 
merely  resembles  that  of  other  dilute  acids. 

3.  Phagocytosis  in  White  Corpuscles. — Mix  some  very  fine  pigment 
granules,  bacterial  emulsion,  or  charcoal  with  a  few  drops  of  frog's  blood, 
let  stand  for  10  or  20  minutes,  then  mount  a  drop  on  the  glass  slide  or 
make  a  smear  and  stain  and  examine  under  a  high- magnifying  microscope. 
Or  inject  a  few  drops  of  one  of  these  pigments  suspended  in  physiological 
saline  and  after  a  few  minutes  examine  drops  of  the  blood  as  above.     In  a 
favorable  field  here  and  there  will  be  found  some  white  corpuscles  which 
have  taken  up  the  pigment.     Colored  corpuscles  have  been  observed  with 
fragments   of   pigment   embedded   in    their   substance.     White    corpuscles 
have  also  been  seen  in  diseased  states  of  the  body  to  contain  micro-organisms, 
for  example  bacilli,  and  have  the  power  of  destroying  these  organisms,  which 
gives  them  the  name  phagocytes. 

4.  Enumeration   of   the   Blood   Corpuscles. — Several    methods   are 
employed  for  counting  the  blood  corpuscles,  most  of  them  depending  upon 
the  same  principle;  i.e.,  the  dilution  of  a  minute  volume  of  blood  with  a 
given  volume  of  a  colorless  solution  similar  in  specific  gravity  to  blood  plasma, 
so  that  the  size  and  shape  of  the  corpuscles  are  altered  as  little  as  possible. 
A  minute  quantity  of  the  well-mixed  solution  is  then  taken,  examined 
under  the  microscope,  either  in  a  flattened  capillary  tube  (Malassez)  or  in 
a  cell  (Hayem  and  Nache,  Cowers)  of  known  capacity,  and  the  number  of 


THE    NUMBER    OF    CORPUSCLES 


159 


corpuscles  in  a  measured  length  of  the  tube  or  in  a  given  area  of  the  cell 
is  counted.  The  length  of  the  tube  and  the  area  of  the  cell  are  ascertained 
by  means  of  a  micrometer  scale  in  the  microscope  ocular;  or,  in  the  case  of 


Depth 

0.100mm 


4OO 


DOME  NH'BAIER  RULING 


FIG.  130. — Section  of  the  new  Bausch  and  Lomb  form  of  the  Thoma-Zeiss  Hemacy- 
tometer.  This  form  has  the  advantage  of  having  the  two  standard  graduated  scales 
reproduced  in  the  next  figure  ground  in  the  glass  slide  itself. 

Gowers's  modification,  by  the  division  of  the  cell  area  into  squares  of  known 
size.  Haying  ascertained  the  number  of  corpuscles  in  the  diluted  blood,  it 
is  easy  to  find  out  the  number  in  a  given  volume  of  normal  blood. 

The  hemacytometer,  which  is  most  used  at  the  present  time,  is  known 
as  the  Thoma-Zeiss  hemacytometer.  It  consists  of  a 
carefully  graduated  pipet,  in  which  the  dilution  of  the 
blood  is  done;  this  is  so  formed  that  the  capillary  stem 
has  a  capacity  equaling  one-hundredth  of  the  bulb  above 
it.  If  the  blood  is  drawn  up  in  the  capillary  tube  to 
the  line  marked  i,  figure  131,  the  saline  solution  may 
afterward  be  drawn  up  the  stem  to  the  line  101 ;  in 
this  way  we  have  101  parts  of  which  the  blood  forms  i. 
As  the  content  of  the  stem  can  be  displaced  unmixed  we 
shall  have  in  the  mixture  the  proper  dilution.  The 
blood  and  the  saline  solution  are  well  mixed  by  shak- 


Neubauer  Ruling  Fuchs-Rosenthal  Ruling 

FIG.  1300. — Standard  hemacytometer  rulings,  1/400  square 
millimeter. 


FIG.  131.— Thoma- 
Zeiss  Hemacytome- 
ter, pipet. 


ing  the  pipet,  in  the  bulb  of  which  is  contained  a  small  glass  bead  for  the 
purpose  of  aiding  the  mixing.  The  counting  instrument  consists  of  a  glass 
slide,  figure  130,  provided  with  a  depressed  area  the  surface  of  which  is 


l6o  THE    BLOOD 

accurately  ruled  so  as  to  present  one  square  millimeter  divided  into  400 
squares  of  one-twentieth  of  a  millimeter  each.  In  the  older  instruments 
the  rulings  are  on  the  cover  disc.  The  micrometer  surface  is  ground 
below  the  general  surface  exactly  one-tenth  millimeter.  If  a  drop  of  the 
diluted  blood  be  placed  upon  the  ruled  surface,  and  covered  with  a  per- 
fectly flat  cover-glass,  the  volume  of  the  diluted  blood  above  each  of 
the  squares  of  the  micrometer,  i.e.,  above  each  3^oo  square  millimeter  area 
will  be  J^ooo  °f  a  cubic  millimeter.  An  average  of  ten  or  more  squares  is 
then  counted,  and  this  number  multiplied  by  4000  X  100  gives  the  number 
of  corpuscles  in  a  cubic  millimeter  of  undiluted  blood.  A  separate  pipet 
is  used  for  making  dilutions  for  counts  of  leucocytes.  In  this,  the  dilution 
is  made  of  one  part  of  blood  and  ten  parts  of  diluting  fluid.  Acetic  acid, 
0.2  of  one  per  cent.,  is  usually  employed  for  this  purpose. 

5.  The  Percentage  of  Corpuscles  and  Plasma  in  Human  Blood.— 
Fill  the  two  graduated  capillary  tubes  of  a  hematocrite  with  blood  drawn 
from  the  tip  of  your  own  finger,  insert  into  the  instrument,  and  centrifuge 
as  rapidly  as  possible.     The  experiment  must  be  performed  within  the  time 
limit  of  clotting  in  order  to  be  successful.     The  corpuscles  will  be  thrown 
down  and  the  percentage  of  plasma  and  corpuscles  can  be  read  off  directly. 
Should  one  fail  to  fill  the  tube  exactly  full,  then  the  percentage  of  plasma 
and  corpuscles  can  be  calculated  from  the  proportion  which  each  bears  to 
the  quantity  in  the  tube. 

6.  Estimation  of  the  Percentage  of  Hemoglobin. — The  percentage 
of  hemoglobin  in  a  sample  of  blood  can  best  be  obtained  by  either  the  Dare 
or  the  Sahli  hemoglobinometer.     The  principle  of  the  Dare  is  given  in  the 
text.     It  rests  on  a  comparison  of  the  color  of  a  drop  of  undiluted  blood 
with  a  standard  color  scale  when  illuminated  by  candle  light.     Sterilize 
a  lobe  of  the  ear  or  finger  tip,  draw  a  drop  of  blood,  bring  the  edge  of  the 
Ware  blood  pipette  carefully  against  the  blood  drop,  allowing  it  to  flow 
between  the  plates.     Place  this  in  the  instrument  and  examine  either  in  a 
dark  room  or  with  the  instrument  turned  toward  a  dark  wall  or  paper. 

A  sample  of  blood  may  be  taken  by  a  Miescher  pipette  diluted  in  the 
Miescher  hemoglobinometer,  and  its  percentage  of  hemoglobin  computed. 
See  text,  page  140,  for  diagram  of  instrument  and  description  of  the 
method. 

Perhaps  a  more  convenient  and  certainly  a  quicker  method  for  deter- 
mining the  percentage  of  hemoglobin  is  Talquist's  hemoglobinometer.  By 
this  method  a  drop  of  blood  is  drawn  directly  on  to  absorbent-paper  furnished 
with  the  instrument,  and  the  resulting  stain  is  compared  directly  with  a  paper 
color  scale  which  is  graduated  in  percentage.  In  this  method  the  comparison 
is  made  in  ordinary  daylight,  and  because  of  its  rapidity  it  is  very  convenient 
for  clinical  examinations,  though  it  is  less  accurate. 


COAGULATION    OF   BLOOD 


161 


7.  Reaction  of  Blood  Plasma. — Wet  a  piece  of  red  litmus-paper  in 
saturated  magnesium  sulphate  solution,  then  touch  one  end  of  the  strip  with 
a  drop  of  blood  drawn  from  your  finger  under  sterile  conditions.     After  a 
few  moments  wash  off  the  excess  of  corpuscles  in  neutral  distilled  water. 
The  blue  at  the  point  of  contact  with  the  blood  indicates  alkalinity. 

8.  The    Specific    Gravity    of    Blood.— From    standard    mixtures    of 
chloroform  and  benzol  with  specific  gravity  of  i .  050,  i .  060,  and  i .  070  make 
up  a  set  of  specific-gravity  solutions  of  1.050,  1.052,  1.054,  etc.,  to  1.070. 
These  standards  may  be  kept  in  stoppered  4-dram  vials,  or  in  test-tubes. 
The  specific  gravity  of  blood  is  determined  by  inserting  with  a  pipet  a 
drop  of  freshly  drawn  blood  into  the  middle  of  one  of  the  solutions,  say 
i  .056.     Since  the  blood  does  not  mix  with  the  chloroform  and  benzol,  the 
drop  will  rise  or  sink  according  to  its  relative  specific  gravity.     By  a  few 
trials  one  may  quickly  find  a  specific  gravity  in  which  the  drop  of  blood 
floats  without  rising  or  sinking.     This  represents  the  specific  gravity  of  the 
drop  of  blood. 

This  method  permits  rapid  clinical  application  and  has  proven  of  con- 
siderable interest  in  the  hands  of  clinicians. 

9.  The  Isotonicity  of  Blood. — The  absorption  or  loss  of  water  by  the 
corpuscles  of  blood  in  solutions  of  other  concentrations  than  that  of 
blood  plasma  can  be  used  as  a 

means  of  determining  the  isoton- 
icity  of  blood.  Make  up  a  series 
of  solutions  of  sodium  chloride, 
varying  by  tenths,  from  0.5  to  1.2 
per  cent.  Prepare  a  series  of 
slides  with  vaselin  rings  and 
mount  drops  of  human  blood  in 
drops  of  saline  of  0.4,  0.6,  0.8,  i, 
1.2  and  1.4  per  cent.,  examine 
immediately  then  every  ten  min- 
utes under  a  high-power  micro- 
scope. The  corpuscles  of  some 
of  the  slides  will  swell  up  and 
may  disintegrate,  others  will 
show  crenation  as  in  figure  126. 
In  the  isotonic  solutions  the  cor- 
puscles will  appear  of  their  nor- 
mal size  and  condition. 

10.  Coagulation  of  Blood. — a.  formal  Clot. — Anesthetize  a  dog,  insert 
a  cannula  into  the  carotid  or  femoral  artery,  and  draw  samples  of  blood 
into  two  or  three  clean,  dry  test  tubes.     Draw  one  sample  into  a  test- 
tube  that  has  had  its  sides  oiled.     Note  the  exact  time  at  which  the  blood 


FIG.  132. — Microscopic  View  of  Clot  Showing 
Fibrin  Network. 


162  THE   BLOOD 

was  drawn  into  the  test-tubes  and  set  the  test-tubes  in  a  test-tube  rack. 
Examine  at  intervals  of  30  seconds  by  gently  inclining  the  test  tubes. 
Presently  it  will  be  noted  that  the  blood  becomes  more  viscous  and  does 
not  flow  freely  up  the  sides  of  the  test-tubes.  Later  the  whole  mass  will 
become  jelly-like  and  will  retain  the  form  of  the  test-tube.  Note  the  time 
of  the  first  slight  change,  and  also  when  the  clot  becomes  more  perfect. 
The  sample  in  the  oiled  test-tube  will  be  found  to  clot  more  slowly. 

If  the  test-tubes  of  clotted  blood  are  left  standing  for  a  day,  the  coagu- 
lum  will  become  smaller  in  size  and  a  transparent  yellowish  blood  will  make 
its  appearance  on  the  surface  or  between  the  sides  of  the  clot  and  the  test- 
tube  wall.  This  fluid  is  the  serum  and  it  is  squeezed  out  by  the  shrinking 
of  the  fibrin  which  holds  the  corpuscles  in  its  meshes. 

b.  The  Time  of  Blood  Clotting. — The  speed  of  clotting  is  measured  more 
accurately  by  Cannon's  coagulometer,  see  figure  108.  A  sample  of  blood  is 
carefully  drawn  from  an  artery  under  conditions  which  insure  fresh  circulat- 
ing blood  (Cannon  and  Mendenhall,  American  Journal  of  Physiology,  Vol.  34, 
p.  225,  for  fuller  details).  This  sample  is  inserted  in  the  coagulometer  and 
successive  tests  for  coagulation  made  every  30  seconds.  Even  a  thread  or  two 
of  fibrin  is  indicated  by  the  apparatus  if  the  lever  is  accurately  counterpoised. 


FIG.  1320. — Successive  tests  of  the  coagulation  of  blood  drawn  from  the  femoral 
artery  of  an  animal  in  uniform  condition.  The  mark  below  the  time  record  signi- 
fies when  the  sample  was  drawn.  Time  in  30  second  intervals.  (From  Cannon  and 
Mendenhall.) 

c.  Microscopic  Examination  of  the  Process  of  Clotting. — Take  a  drop  of 
fresh  blood  from  the  tip  of  your  finger  under  sterile  conditions  and  mount 
on  a  microscopic  slide  in  a  few  drops  of  salt  solution,  and  examine  immediately 
under  the  high  power.     Small  threads  of  fibrin  will  presently  be  seen  to  form 
across  the  field,  usually  being  most  clearly  obvious  where  fragments  of 
white  corpuscles  are  noted,  see  figures  107  and  132.     The  threads  of  fibrin 
become  more  apparent  when  stained  with  rosanilin. 

d.  Whipped  Blood.— Draw  a  sample  of  blood  into  a  glass  tumbler, 
enough  to  fill  it  one-half  or  two-thirds  full.     Immediately  begin  vigorously 
stirring  the  blood  with  a  bunch  of  stiff  wires  or  a  pencil,  and  keep  it  up  until 
the  time  of  clotting  has  passed,  5  or  10  minutes.     In  this  instance  the  wires 
will  break  up  and  collect  the  fibrin  as  fast  as  it  forms,  and  no  firm  mass  will 
be  produced.     The  remaining  fluid  is  called  whipped  blood.     The  fibrin  can 
be  removed  from  the  wires  and  washed  in  tap  water  until  all  the  adherent 
red  corpuscles  are  removed.     This  mass  of  fibrin  is  white,  elastic,  and  com- 
posed of  a  network  of  thread-like  fibers.     It  is  these  fibers  extending 


CHEMISTRY    OF   BLOOD  163 

through  and  through  the  mass  of  blood  which  makes  it  retain  the  form  of 
the  vessel  when  undisturbed  clotting  occurs. 

e.  The  Influence  of  Salt  Solution  on  Blood  Clotting. — Add  2  cc.of  saturated 
magnesium  sulphate,  i  per  cent,  sodium  oxalate,  and  0.9  per  cent,  sodium 
chloride  to  each  of  three  test-tubes.  Draw  into  each  test-tube  5  to  6  cc.  of 
blood  and  immediately  mix  thoroughly  and  let  stand.  The  magnesium 
and  oxalate  test-tubes  will  not  coagulate  even  though  -they  stand  for  days, 
but  the  sodium-chloride  blood  will  clot  in  a  few  minutes. 

The  magnesium-sulphate  blood  will  coagulate  if  diluted  with  a  sufficient 
amount  of  distilled  water  or  physiological  saline  solution.  Make  a  series 
of  dilutions  and  note  when  coagulation  takes  place.  The  sodium-oxalate 
blood  will  coagulate  when  a  sufficient  excess  (of  i  per  cent,  solution)  of 
calcium  chloride  is  carefully  added  to  neutralize  the  excess  of  sodium 
oxalate.  Demonstrate  these  on  a  series  of  samples. 

If  a  quantity  of  magnesium  or  oxalate  blood  is  secured  and  separated  by 
a  centrifuge  or  by  letting  stand  for  a  sufficient  time,  a  sample  of  uncoagulated 
plasma  will  be  obtained.  This  sample  will  coagulate  when  it  is  treated  as 
just  described  above  for  blood,  showing  that  the  antecedents  of  fibrin  are 
found  in  the  plasma. 

/.  Action  of  Tissue  Extracts  on  Coagulation. — Wash  out  the  blood  of 
a  small  animal  by  circulating  0.9  per  cent,  saline  through  the  arteries  until 
the  outflowing  fluid  from  the  veins  is  clear.  Take  an  organ,  the  liver  for 
example,  grind  it  up  in  a  sausage  mill  by  running  it  through  the  mill  two 
or  three  times,  then  extract  with  0.9  per  cent,  physiological  saline.  The 
macerating  mass  should  be  shaken  up  at  intervals,  and  may  be  kept  from 
spoiling  by  adding  an  excess  of  chloroform  or  by  keeping  on  ice.  A  few 
cubic  centimeters  of  this  fluid  extract  added  to  a  sample  of  freshly  drawn 
blood  will  very  greatly  hasten  the  rapidity  of  coagulation.  This  tissue  ex- 
tract contains  the  thrombqplastin  of  Howell  (Thrombokinase  of  Morawitz), 
and  hastens  the  formation  of  thrombin  from  thrombogen. 

ii.  The  Chemistry  of  Blood  Plasma  (or  Serum). — The  blood  plasma 
contains  all  the  chemical  substances  which  are  utilized  by  the  tissues  in 
their  nutrition  or  which  are  thrown  off  by  the  tissues  as  a  result  of  their 
activity.  It  is  therefore  a  very  complex  mixture.  The  serum  contains  the 
same  substances  in  the  same  proportion,  with  the  exception  of  the  antece- 
dents of  fibrin.  It  may,  therefore,  be  used  as  a  substitute  for  plasma  in 
most  cases. 

a.  Proteins  of  Plasma. — There  are  three  principal  proteins  in  blood 
plasma:  serum-albumin,  serum-globulin,  and  fibrinogen.  These  may  be 
isolated  as  follows:  To  a  sample  of  blood  plasma  add  an  equal  quantity  of 
sodium-chloride  solution  that  has  been  saturated  at  40°  C.  A  white  floccu- 
lent  precipitate  of  fibrinogen  comes  down.  Filter  off,  and  add  to  the  filtrate 


1 64  THE   BLOOD 

an  equal  volume  of  saturated  ammonium  sulphate.  A  second  heavier  pre- 
cipitate of  serum- globulin  separates  out.  When  this  is  separated,  and  crys- 
tals of  ammonium  sulphate  are  added  to  the  filtrate  to  complete  saturation 
at  40°  C.,  a  third  precipitate  of  serum-albumin  separates. 

Each  of  these  precipitates  may  be  redissolved  and  purified  by  reprecipi- 
tation  and  can  be  tested  by  the  characteristic  protein  reactions,  see  page 
107,  which  they  all  give. 

b.  Sugars  of  Blood  Plasma  or  Serum. — If  a  quantity  of  blood  serum  is 
diluted  with  about  5  to  10  times  its  volume  of  water,  and  the  proteins  are 
removed  by  slight  acidulation  with  acetic  acid  and  boiling  and  filtering,  the 
filtrate  will  contain  reducing  sugar  and  the  various  salts  of  blood  plasma. 
To  a  concentrated  sample  of  the  filtrate  add  Fehling's  solution  and  boil. 
A  reddish  precipitate  indicates  the  presence  of  reducing  sugar.     If  this  ex- 
periment is  done  quantitatively,  about  from  o.i  to  0.2  percent,  of  sugar  will 
be  found.     The  sugar  may  be  separated  from  the  serum  by  dialysis  through 
collodian  membranes. 

c.  The  Salts  of  Blood  Plasma. — The  salts  of  blood  plasma  are  tested 
best  by  evaporating  some  of  the  blood  serum  to  dryness,  and  burning  the 
residue  to  oxidize  the  organic  matter  and  dissolving  the  ash  in  water.     Test 
as  follows:  To  a  sample  add  i  per  cent,  of  silver  nitrate;  a  white  precipitate 
soluble  in  an  excess  of  ammonia,  but  not  soluble  in  nitric  acid,  indicates 
chlorides. 

To  a  second  sample  add  i  per  cent,  barium  chloride.  If  sulphates  are 
present  there  will  be  a  white  precipitate  which  settles  out  quickly. 

Acidify  a  third  sample  with  nitric  acid  and  add  ammonium  molybdate 
and  heat.  A  yellow  precipitate  indicates  the  presence  of  phosphates. 

To  the  fourth  sample  add  an  excess  of  strong  ammonia  and  i  per 
cent,  ammonium  oxalate,  heat.  A  white  precipitate  indicates  the  presence 
of  calcium. 

12.  Blood  Corpuscles. — The  characteristic  substance  in  the  composi- 
tion of  the  blood  corpuscles  is  the  pigment  known  as  hemoglobin,  and  this  is 
the  only  chemical  factor  that  will  be  considered  in  these  experiments. 

a.  Hemoglobin  Crystals. — Take  a  sample  of  dog's  blood,  or  if  a  centri- 
fuge is  available  separate  and  wash  a  sample  of  blood  corpuscles,  and 
mix  with  about  three  volumes  of  saturated  ether  water,  or  if  blood  is  used 
dilute  with  two  or  three  volumes  of  water  and  add  about  10  per  cent,  by 
volume  of  pure  ether  and  shake  thoroughly.     Crystals  of  oxyhemoglobin 
will  be  formed,  and  these  can  be  mounted  and  examined  with  a  microscope. 

b.  Spectrum  of  Hemoglobin  and  its  Compounds. 

i.  Oxyhemoglobin. — Dilute  a  sample  of  defibrinated  blood  with  about 
ten  volumes  of  distilled  water.  From  this  stock  solution  make  five  solutions 
all  differing  by  33^  per  cent.  Examine  these  with  a  direct-vision  spectro- 
scope. Make  a  drawing  showing  the  absorption  spectrum  of  each  sample 


HEMOGLOBIN  165 

as  compared  with  the  solar  spectrum.     Compared  with  the  spectrum 
shown  in  the  frontispiece. 

2.  Hemoglobin. — The  oxygen  can  be  driven  out  from  the  oxyhemoglobin 
by  adding  to  the  above  samples  a  few  drops  of  ammonium  sulphide  and 
gently  warming.     Re-examine  with  the  direct- vision  spectroscope  and  map 
as  before. 

3.  Carbon-monoxide  Hemoglobin. — Pass  a  stream  of  ordinary  illumi- 
nating gas  through  the  dilutions  of  hemoglobin.     The  carbon  monoxide 
of  the  gas  will  form  a  compound  with  the  hemoglobin,  which  now  turns  a 
bright  scarlet  color.     When  examined  with  the  spectroscope,  the  absorp- 
tion bands  are  found  to  be  very  similar  to  those  of  oxyhemoglobin.     How- 
ever, map  the  spectrum  to  the  scale  as  usual.     Add  the  reducing  agent, 
warm,  and  shake  vigorously  and  re-examine.     It  is  very  difficult  to  break  up 
the  combination  of  hemoglobin  with  carbon  monoxide,  hence  the  poisonous 
action  of  this  gas. 


CHAPTER  V. 
THE  CIRCULATION  OF  THE  BLOOD 

THE  blood  is  contained  in  a  system  of  closed  vessels  through  which  it  is 
kept  in  circulation  during  the  life  of  the  individual.  The  energy  to  keep  up 
this  motion  is  supplied  by  the  heart,  which  is  a  large  muscular  organ  con- 
sisting of  four  great  divisions,  the  right  and  left  auricles  and  right  and  left 
ventricles.  The  right  ventricle  discharges  its  blood  into  the  pulmonary 


FIG.  133. — Diagram  of  the  Circulation  in  an  Animal  with  a  Completely  Separated 
Right  and  Left  Ventricle  and  a  Double  Circulation.  Ad,  Right  auricle  receiving  the 
superior  and  inferior  venae  cavae,  Vcs  and  Vci;  Dth,  thoracic  duct,  the  main  trunk  of  the 
lymphatic  system;  Ad,  right  auricle;  Vd,  right  ventricle;  Ap,  pulmonary  artery;  P,  lung; 
Vp,  pulmonary  vein;  As,  left  auricle;  Vs,  left  ventricle;  Ao,  aorta;  D,  intestine;  L,  liver; 
Vp,  portal  vein;Z,-y,  hepatic  vein.  (After  Huxley.) 

artery,  through  which  it  passes  to  the  lungs,  returning  through  the  pulmonary 
veins  to  the  left  auricle,  and  into  the  ventricle.  From  the  left  ventricle 
the  blood  is  pumped  into  the  great  aorta,  and  through  its  branches  distrib- 
uted to  the  entire  body.  The  terminal  arteries  are  continuous  with  the 
IT  166 


THE    HEART  167 

general  capillaries  of  the  body,  and  these  in  turn  with  the  veins,  which  con- 
duct the  blood  back  to  the  right  side  of  the  heart  again.  It  will  be  seen, 
therefore,  that  the  circulatory  apparatus  consists  of  two  great  divisions,  the 
pulmonary  and  the  systemic  circulation.  This  arrangement  is  illustrated 
by  the  accompanying  figure.  A  study  of  this  figure  will  show  that  in  certain 
regions  of  the  systemic  circulation  there  are  two  capillary  beds  between  the 
main  arteries  and  the  main  veins.  This  subordinate  stream  through  the 
liver  is  called  the  portal  circulation,  and  the  similar  arrangement  existing 
in  the  kidney  is  called  the  renal  circulation.  This,  in  general,  is  the  outline 
of  the  course  of  the  blood  in  its  circulation. 

To  make  a  study  of  the  various  phenomena  manifested  in  the  physiology 
of  the  circulatory  apparatus,  it  is  obvious  that  we  have  to  do  with  certain 
fundamental  activities;  first,  the  physiology  of  the  pumping  organ,  the  heart; 
second,  the  movement  of  the  blood  in  the  arteries,  capillaries,  and  veins; 
third,  the  co-ordination  of  these  various  divisions  of  the  apparatus  through 
the  nervous  system.  To  understand  this  it  will  be  necessary  to  have  in 
mind  in  detail  the  anatomical  structure  of  the  apparatus  itself. 

ANATOMICAL  CONSIDERATIONS. 

The  Heart. — The  heart  is  contained  in  the  chest  or  thorax,  and  lies 
between  the  right  and  left  lungs,  figure  134,  enclosed  in  a  membranous  sac, 
the  pericardium.  The  pericardium  is  made  up  of  two  distinct  parts,  an 
external  fibrous  membrane  and  an  internal  serous  layer  which  not  only 
lines  the  fibrous  sac,  but  also  is  reflected  on  to  the  heart,  which  it  completely 
invests.  These  form  a  closed  sac,  the  cavity  of  which  contains  just  enough 
pericardial  fluid  to  lubricate  the  two  surfaces,  and  thus  to  enable  them  to  glide 
smoothly  over  e'ach  other  during  the  movements  of  the  heart.  The  vessels 
passing  in  and  out  of  the  heart  receive  investments  from  this  sac  to  a  greater 
or  less  degree. 

The  heart  is  situated  in  the  chest  behind  the  sternum  and  costal  carti- 
lages, being  placed  obliquely  from  right  to  left.  It  is  of  pyramidal  shape, 
with  the  apex  pointing  downward,  outward,  and  toward  the  left,  and  the 
base  backward,  inward,  and  toward  the  right.  The  heart  is  suspended  in 
the  chest  by  the  large  vessels  which  proceed  from  its  base,  but,  excepting 
at  the  base,  the  organ  itself  hangs  free  within  the  sac  of  the  pericardium. 
The  part  which  rests  upon  the  diaphragm  is  flattened,  and  is  known  as  the 
diaphragmatic  surface,  while  the  free  upper  part  is  called  the  sternocostal 
surface. 

On  examination  of  the  external  surface  the  division  of  the  heart  into  parts 
which  correspond  to  the  chambers  inside  of  it  may  be  traced.  A  deep  trans- 
verse groove,  called  the  coronary  sulcus,  divides  the  auricles  from  the  ventri- 
cles; and  the  anterior  longitudinal  sulcus  runs  between  the  ventricles,  both  in 


1 68 


THE    CIRCULATION    OF    THE   BLOOD 


front  and  in  the  back,  separating  the  one  from  the  other.  The  anterior  groove 
is  nearer  the  left  margin,  and  the  posterior  nearer  the  right,  as  the  front 
surface  of  the  heart  is  made  up  chiefly  of  the  right  ventricle  and  the  posterior 
surface  of  the  left  ventricle.  The  coronary  vessels  which  supply  the  tissue 
of  the  heart  with  blood  run  in  the  furrows  or  sulci;  also  the  nerves  and  lymph- 
atics, which  are  embedded  in  more  or  less  fatty  material,  are  found  in  this 
groove. 

The  Chambers  of  the  Heart. — The  interior  of  the  heart  is  divided  by  a 
longitudinal  partition  in  such  a  manner  as  to  form  two  chief  chambers  or 
cavities,  the  right  and  the  left.  Each  of  these  chambers  is  again  subdivided 
transversely  into  an  upper  and  a  lower  portion,  called,  respectively,  the  auricle 


FIG.  134. — Outline  of  Heart,  Lungs,  and  Liver  to  Show  their  Relations  to  each  other  and  to 
the  Chest  Wall.     (Heusman  and  Fisher's  "Anatomical  Outlines.") 

and  the  ventricle,  which  freely  communicate.  The  aperture  of  communica- 
tion, however,  is  guarded  by  valves  so  disposed  as  to  allow  blood  to  pass 
freely  from  the  auricle  into  the  ventricle,  but  not  in  the  opposite  direction. 
There  are  thus  four  cavities  in  the  heart,  the  auricle  and  ventricle  of  one  side 
being  quite  separate  from  those  on  the  other,  figure  135. 

The  right  auricle,  the  right  part  of  the  base  of  the  heart  as  viewed  from 
the  front,  is  a  thin-walled  cavity  of  more  or  less  quadrilateral  shape,  pro- 
longed at  one  corner  into  a  tongue-shaped  portion,  the  right  auricular  appen- 
dix, which  slightly  overlaps  the  exit  of  the  aorta  from  the  left  ventricle. 

The  interior  of  the  auricle  is  smooth,  being  lined  with  the  general  lining 
membrane  of  the  heart,  the  endocardium.  The  superior  and  inferior  vena 
cavce  open  into  the  auricle.  The  opening  of  the  inferior  cava  is  protected 


THE   HEART 


1 69 


and  partly  covered  by  a  membrane  called  the  Eustachian  valve.  In  the 
posterior  wall  of  the  auricle  is  a  slight  depression  called  the  fossa  ovalis, 
which  corresponds  to  an  opening  between  the  right  and  left  auricles,  exist- 
ing in  fetal  life  the  foramen  ovale.  The  foramen  fails  to  close  in  many 
individuals.  Statistics  from  observations  of  hearts  from  the  dissecting 
rooms  show  as  many  as  forty  out  of  a  hundred  hearts  with  more  or  less 
open  interauricular  foramina.  In  the  appendix  are  closely  set  elevations 
of  the  muscular  tissue  covered  with  endocardium,  and  on  the  anterior 


FIG.  135. — The  Right  Auricle  and  Ventricle  Opened  and  a  Part  of  their  Right  and 
Anterior  Walls  Removed  so  as  to  Show  their  Interior,  i,  Superior  vena  cava;  2,  inferior 
vena  cava;  2',  hepatic  veins  cut  short;  3,  right  auricle;  3',  placed  in  the  fossa  ovalis,  below 
which  is  the  Eustachian  valve;  3",  is  placed  close  to  the  aperture  of  the  coronary  vein; 
f,  t>  placed  in  the  auriculo-ventricular  groove,  where  a  narrow  portion  of  the  adjacent 
walls  of  the  auricle  and  ventricle  has  been  preserved;  4,  4,  cavity  of  the  right  ventricle, 
the  upper  figure  is  immediately  below  the  semilunar  valves;  4',  large  columna  carnea  or 
musculus  papillaris;  5,  5',  5",  tricuspid  valve;  6,  placed  in  the  interior  of  the  pulmonary 
artery,  a  part  of  the  anterior  wall  of  that  vessel  having  been  removed  and  a  narrow  portion 
of  it  preserved  at  its  commencement  where  the  semilunar  valves  are  attached;  7,  concavity 
of  the  aortic  arch,  close  to  the  cord  of  the  ductus  arteriosus;  8,  ascending  part  or  sinus  of 
the  arch  covered  at  its  commencement  by  the  auricular  appendix  and  pulmonary  artery;  9, 
placed  between  the  innominate  and  left  carotid  arteries;  10,  appendix  of  the  left  auricle; 
n,  II,  outside  of  the  left  ventricle  the  lower  figure  near  the  apex.  (Allen  Thomson.) 

wall  of  the  auricle  are  similar  elevations  arranged  parallel  to  one  another, 
called  musculi  pectinati. 


170  THE    CIRCULATION    OF   THE   BLOOD 

The  right  ventricle  forms  the  right  margin  of  the  heart.  It  takes  no 
part  in  the  formation  of  the  apex.  On  section  its  cavity  is  semilunar  or 
crescentic,  figure  135.  Into  it  are  two  openings,  the  venous  orifice  at  the 
base,  and  the  arterial  orifice  of  the  pulmonary  artery,  also  at  the  base  but 
more  to  the  left.  The  part  of  the  ventricle  leading  to  the  pulmonary  artery 
is  called  the  conus  arteriosus.  Both  orifices  are  guarded  by  valves,  the 
former  called  the  tricuspid  and  the  latter  the  semilunar.  In  this  ventricle 
are  also  the  projections  of  the  muscular  tissue  called  the  trabecula  carnece. 

The  left  auricle  is  situated  at  the  left  and  posterior  part  of  the  base  of 
the  heart.  The  left  auricle  is  only  slightly  thicker  than  the  right  and  its 
form  and  structure  are  the  same  as  in  the  right.  The  left  venous  orifices 
are  oval  and  a  little  smaller  than  those  on  the  right  side  of  the  heart.  There 
is  a  slight  vestige  of  the  foramen  on  the  septum  between  the  auricles. 


FIG.  136. — Cross-section  of  a  Completely  Contracted  Human  Heart,  at  the  Level  of  the 
Lower  and  Middle  Thirds.     (According  to  Krehl.) 

The  left  ventricle  occupies  the  posterior  and  apical  portion  of  the  heart, 
and  is  connected  directly  with  the  great  aorta.  It  is  separated  from  the 
auricle  by  the  bicuspid  or  mitral  valve,  and  the  opening  into  the  great  aorta 
is  guarded  by  the  semilunar  valves.  The  walls  of  the  left  ventricle  are  two 
or  three  times  as  heavy  as  those  of  the  right,  and  may  be  as  much  as  half  an 
inch  in  total  thickness. 

The  left  ventricle  is  capable  of  containing  90  to  120  c.c.  of  blood.  The 
capacity  of  the  auricles  is  considerably  less  after  death  owing  to  their  con- 
tracted condition.  The  whole  heart  is  about  12  cm.  long  by  8  cm.  at  its 
greatest  width,  and  6  cm.  in  thickness.  The  average  weight  in  the  adult  is 
about  300  grams. 

The  walls  of  the  heart  are  constructed  almost  entirely  of  layers  of  muscu- 
lar fibers;  but  a  ring  of  connective  tissue,  to  which  some  of  the  muscular 


THE    HEART 


171 


fibers  are  attached,  is  inserted  between  each  auricle  and  ventricle  and  forms 
the  boundary  of  the  venous  opening.  Fibrous  tissue  also  exists  at  the 
origins  of  the  pulmonary  artery  and  aorta.  The  muscular  fibers  of  each 
auricle  are  in  part  continuous  with  those  of  the  others  and  in  part  separate; 
and  the  same  holds  true  for  the  ventricles.  The  fibers  of  the  auricles  are, 


FIG.  137.  FIG.  138. 

FIG.  137. — Cardiac  Muscle  Cells,  Showing  their  Form,  Branches,  Nuclei,  and  Striae. 
From  the  heart  of  a  young  rabbit.  Magnified  425  diameters,  a,  Line  of  junction  between 
the  cells  (intercellular  cement) ;  b,  c,  branches  of  the  cells.  (Schafer.) 

FIG.  138.— Cardiac  Muscle  Cells  of  the  Left  Ventricle  of  a  Child  at  Birth  (full  term),  to 
show  the  form  of  the  cells,  their  structural  details,  their  relations  to  one  another,  and  their 
general  agreement  with  those  of  cold-blooded  vertebrates.  /4,  Large  cell  with  two  nuclei; 
this  cell  has  nearly  the  proportions  of  those  of  the  adult;  B,  group  of  cells  in  their  natural 
relation.  At  the  right  of  the  middle  cell  are  two  spaces  or  fissures,  n,  Nucleus.  The 
transverse  striations  cross  the  nuclei  in  all  the  cells,  and  each  nucleus  possesses  several 
nucleoli.  (Gage.) 

however,  quite  separate  from  those  of  the  ventricles.  The  bond  of  con- 
nection between  the  auricles  and  the  ventricles  is  made  by  the  Purkinje 
fibers,  an  embryonic  muscular  type  of  tissue  composing  the  auriculo- 
ventricular  strand  in  the  septum  called  the  bundle  of  His. 

The  development  of  the  heart  shows  that  it  is  derived  from  an  embry- 
onic tube,  which  in  its  growth  becomes  twisted  upon  itself  and  divided  into 
the  two  main  divisions  that  we  know  in  the  adult.  Anatomical  dissections 
have  shown  that  the  muscles  of  the  ventricles  form  spiral  sheaths  extending 
from  the  base  of  the  two  ventricles  in  spiral  bands  toward  the  apex.  These 
bands  of  muscle  are  wound  about  the  surface  of  the  ventricles  in  the  right- 


172 


THE    CIRCULATION    OF    THE   BLOOD 


to-left  direction.  At  the  apex  they  extend  up  into  the  deeper  tissue.  If 
the  superficial  muscles  are  dissected  off,  there  is  left  a  great  central  core  of 
muscle,  which  is  described  by  MacCallum  as  running  more  transversely 


* 


FIG.  139. 


FIG.  140. 


FIG.  139. — Diagram  of  the  Course  of  the  Superficial  Muscle  Layers  Originating  in  the 
Right  and  Left  Coronary  Sulci  and  in  the  Posterior  Half  of  the  Tendon  of  the  Conus. 
C,  Anterior  papillary  muscle.  (After  MacCallum.) 

FIG.  140. — Diagram  of  the  Course  of  the  Superficial  Muscle  Layers  Originating  in  the 
Anterior  Half  of  the  Tendon  of  the  Conus.  A,  Posterior  papillary  muscle;  B,  papillary 
muscle  of  the  septum.  (After  MacCallum.) 


FIG.  141. 


FIG.  142. 


FIG.  141. — Diagram  of  the  Course  of  the  Layer  Superficial  to  the  Deepest  Layer  of  the 
Muscle  of  the  Left  Ventricle,  which  is  shown  in  outline.  The  deepest  layer  is  also  shown. 
A,  Posterior  papillary  muscle;  B,  papillary  muscle  of  the  septum.  (After  MacCallum.) 

FIG.  142. — Diagram  of  a  Layer  still  more  Superficial  to  that  Shown  in  Fig.  141,  and 
Ending  in  the  Anterior  Papillary  Muscle.  The  deeper  layers  are  represented  in  dotted 
lines.  A,  Posterior  papillary  muscle;  B,  papillary  muscle  of  septum;  C,  anterior  papillary 
muscle.  (After  MacCallum.) 

around  the  wall  of  one  ventricle,  then  through  the  septum  and  around  the 
other  in  a  reverse  scroll,  figure  141. 

The  Valves  of  the  Heart. — The  valves  of  the  heart  are  arranged  so 
that  the  blood  can  pass  only  in  one  direction.  These  are  the  tricuspid 


THE    ARTERIES  173 

valve,  between  the  right  auricle  and  right  ventricle,  figure  135,  and  the  semi- 
lunar  valve  of  the  pulmonary  artery,  the  mitral  valve  between  the  left  auricle 
and  ventricle,  and  semilunar  valve  of  the  aorta.  The  bases  of  the  tricuspid, 
figure  152,  and  mitral  valves  are  attached  to  the  walls  of  the  venous  orifices 
respectively.  Their  ventricular  surfaces  and  borders  are  fastened  by  slender 
tendinous  fibers,  the  chorda  tendinece,  to  the  internal  surface  of  the  walls 
of  the  ventricles  at  points  which  project  into  the  ventricular  cavity  in  the  form 
of  bundles  or  columns,  the  musculi  papillares. 

The  semilunar  valves  guard  the  orifices  of  the  pulmonary  artery  and  of 
the  aorta.  They  are  nearly  alike  on  both  sides  of  the  heart,  but  the  aortic 
valve  is  altogether  thicker.  Each  valve  consists  of  three  parts  which  are 
of  similunar  shape,  the  convex  margin  of  each  being  attached  to  a  fibrous 
ring  at  the  place  of  junction  of  the  artery  to  the  ventricle,  and  the  concave 
or  nearly  straight  border  being  free,  so  as  to  form  a  little  pouch  like  a  pocket, 
figure  151.  In  the  center  of  each  free  edge  of  the  valves  which  contains 
a  fine  cord  of  fibrous  tissue  is  a  small  fibrous  nodule,  the  corpus  Arantii  of 
the  valves. 

The  Arteries. — The  arterial  system  begins  at  the  left  ventricle  in  a 
single  large  trunk,  the  aorta,  which,  almost  immediately  after  its  origin, 
gives  off  in  the  thorax  three  large  branches  for  the  supply  of  the  head,  neck, 
and  upper  extremities;  it  then  traverses  the  thorax  and  abdomen,  giving 
off  branches,  some  large  and  some  small,  for  the  supply  of  the  various  organs 
and  tissues  it  passes  on  its  way.  In  the  abdomen  it  divides  into  two  chief 
branches.  The  arterial  branches,  wherever  given  off,  divide  and  subdivide 
until  the  caliber  of  each  subdivision  becomes  very  minute.  These  smallest 
arteries  are  called  arterioles.  These  arterioles  are  continuous  into  the  capil- 
laries. Arteries  frequently  communicate  or  anastomose  with  other  arteries. 
The  arterial  branches  are  usually  given  off  at  an  acute  angle,  and  the  areas 
of  the  branches  of  an  artery  generally  exceed  that  of  the  parent  trunk,  and, 
as  the  distance  from  the  origin  is  increased,  the  area  of  the  combined 
branches  is  increased  also.  As  regards  the  arterial  system  of  the  lungs,  the 
pulmonary  artery  and  its  subdivisions,  they  are  branched  in  much  the  same 
manner  as  the  arteries  belonging  to  the  general  systemic  circulation. 

The  walls  of  the  arteries  are  composed  of  three  principal  coats,  the  ex- 
ternal, or  tunica  adventitia,  the  middle,  or  tunica  media,  and  the  internal,  or 
tunica  intima.  The  external  coat,  figures  143  and  144,  a,  the  strongest  and 
toughest  part  of  the  wall  of  the  artery,  is  formed  of  areolar  tissue,  with  which 
is  mingled  throughout  a  network  of  elastic  fibers.  The  middle  coat,  figure 
144,  m,  is  composed  of  both  muscular  and  elastic  fibers  with  a  certain  pro- 
portion or  areolar  tissue.  In  the  larger  arteries,  its  thickness  is  compara- 
tively as  well  as  absolutely  much  greater  than  in  the  small  arteries,  consti- 
tuting, as  it  does,  the  greater  part  of  the  arterial  wall.  The  muscular 
fibers  are  unstriped,  figure  145,  and  are  arranged,  for  the  most  part,  trans- 


THE    CIRCULATION    OF    THE   BLOOD 


versely  to  the  long  axis  of  the  artery,  figure  143,  w,  while  the  elastic  element, 
taking  also  a  transverse  direction,  is  disposed  in  the  form  of  closely  inter- 
woven and  branching  fibers  intersecting  in  all  parts  the  layers  of  muscular 
fiber.  In  arteries  of  various  size  there  is  a  difference  in  the  proportion  of 
the  muscular  and  elastic  element,  elastic  tissue  preponderating  in  the  largest 
arteries  and  unstriped  muscle  in  those  of  medium  and  small  size.  The 
arteries  are  quite  elastic  in  both  large  and  small  vessels.  The  internal  coat 
is  formed  by  a  layer  of  elastic  tissue,  called  thefenestrated  membrane  of  Henle. 
It  is  peculiar  in  its  tendency  to  curl  up  when  peeled  off  from  the  artery,  and 


FIG.  143. 


FIG.  144. 


FIG.  145. 


FIG.  143. — Minute  Artery  Viewed  in  Longitudinal  Section,  e,  Nucleated  endothelial 
membrane,  with  faint  nuclei  in  lumen,  looked  at  from  above;  i,  thin  elastic  tunica  intima; 
m,  muscular  coat  or  tunica  media;  a,  tunica  adventitia.  (Klein  and  Noble  Smith.) 

FIG.  144. — Transverse  Section  through  a  Large  Branch  of  the  Inferior  Mesenteric 
Artery  of  a  Pig.  e,  Endothelial  membrane;  i,  tunica  elastica  interna,  no  subendothelial 
layer  is  seen;  m,  muscular  tunica  media,  containing  only  a  few  wavy  elastic  fibers;  e,  c, 
tunica  elastica  externa,  dividing  the  media  from  the  connective-tissue  adventitia,  a.  Mag- 
nification, 350  diameters.  (Klein  and  Noble  Smith.) 

FIG.  145. — Muscular  Fiber  Cells  from  Human  Arteries.  Magnified  350  diameters, 
a,  Nucleus;  B,  a  fiber  cell  treated  with  acetic  acid.  (Kolliker.) 

in  the  perforated  and  streaked  appearance  which  it  presents  under  the  micro- 
scope. The  inner  surface  of  the  artery  is  lined  with  a  delicate  layer  of  elon- 
gated endothelial  cells  which  make  it  smooth  and  polished  and  furnish  a 
nearly  impermeable  surface  along  which  the  blood  may  flow  with  the 
smallest  possible  amount  of  resistance  from  friction. 

Many  of  the  arteries  are  accompanied  by  a  plexus  of  vaso-motor  nerves. 
In  the  smaller  arteries  these  nerves  consist  of  few  fibers  that  form  a  delicate 
network  over  the  walls  of  the  vessels.  Many  fibers  appear  to  end  in  the 
muscle  cells  of  the  arterioles  in  the  proximity  of  the  nuclei. 

The  Capillaries. — In  all  vascular  textures,  except  some  parts  of  the 
corpora  cavernosa  of  the  penis,  of  the  uterine  placenta,  and  of  the  spleen, 


THE    CAPILLARIES  175 

the  transmission  of  the  blood  from  the  minute  branches  of  the  arteries  to  the 
minute  veins  is  effected  through  a  network  of  capillaries.  They  may  be 
seen  in  all  minutely  injected  preparations. 

The  point  at  which  the  arteries  terminate  and  the  capillaries  commence 
cannot  be  exactly  defined,  for  the  transition  is  gradual.  The  capillaries 
maintain  essentially  the  same  diameter  throughout.  The  meshes  of  the 


FIG.  146. — Vein  and  Capillaries.     Silver-nitrate  and  hematoxylin  stain,  to  show  outlines 
of  endothelial  cells  and  their  nuclei.     (Bailey.) 


network  that  they  compose  are  more  uniform  in  shape  and  size  than  those 
formed  by  the  anastomoses  of  the  minute  arteries  and  veins. 

The  walls  of  the  capillaries  are  composed  of  a  single  layer  of  elongated 
or  radiate,  flattened  and  nucleated  endothelial  cells,  so  joined  and  dove- 
tailed together  as  to  form  a  continuous  transparent  membrane,  figure  146. 


FIG.  147. — Network  of  Capillary  Vessels  of  the  Air  Cells  of  the  Horse's  Lung 
Magnified,  a,  a,  Capillaries  proceeding  from  b,  b,  terminal  branches  of  the  pulmonary 
artery.  (Frey.) 


Outside  these  cells  in  the  larger  capillaries  there  is  a  structureless  supporting 
membrane  on  the  inner  surface  of  which  they  form  a  lining. 

The  diameter  of  the  capillary  vessels  varies  somewhat  in  the  different 
textures  of  the  body,  the  most  common  size  being  about  12  micro  millimeters, 
of  an  inch.     Among  the  smallest  may  be  mentioned  those  of  the 


1 76 


THE    CIRCULATION   OF   THE   BLOOD 


brain  and  of  the  follicles  of  the  mucous  membrane  of  the  intestines;  among 
the  largest,  those  of  the  skin  and  especially  those  of  the  medulla  of  the  bones. 

The  form  of  the  capillary  network  differs  in  the  different  organs  of  the 
body,  but  is  usually  adjusted  to  the  structural  arrangement  of  the  cells  of 
any  given  organ. 

The  capillary  network  is  closest  in  the  lungs  and  in  the  choroid  coat  of 
the  eye.  In  the  human  liver  the  interspaces  are  of  the  same  size,  or  even 


PIG.  148. — Capillaries  of  Striated  Muscular  Tissue.     From  a  cat.     Magnified  300  diam- 
eters.    A,  Artery;  V,  vein.     (Heitzmann.) 


smaller  than  the  capillary  vessels  themselves.  In  the  human  lung  the  spaces 
are  smaller  than  the  vessels;  in  the  human  kidney  and  in  the  kidney  of  the 
dog  the  diameter  of  the  injected  capillaries,  compared  with  that  of  the  inter- 
spaces, is  in  the  proportion  of  one  to  four,  or  one  to  three.  The  brain 
receives  a  very  large  quantity  of  blood;  but  its  capillaries  are  very  minute 
and  are  less  numerous  than  in  some  other  parts.  In  the  mucous  mem- 


THE   VEINS 


177 


branes,  for  example  in  the  conjunctiva  and  in  the  cutis  vera,  the  capillary 
vessels  are  much  larger  than  in  the  brain  and  the  interspaces  narrower, 
namely,  not  more  than  three  or  four  times  wider  than  the  vessels.  In  the 
periosteum  and  in  the  external  coat  of  arteries  the  meshes  are  much  larger, 
their  width  being  about  ten  times  that  of  the  vessels.  It  may  be  held  as  a 
general  rule  that  the  more  active  the  functions  of  an  organ  are,  the  more 
vascular  it  is. 

The  Veins. — The  venous  system  begins  in  small  vessels  which   are 
slightly  larger  than  the  capillaries  from  which  they  spring.     These  vessels 


FIG.  149. — Transverse  Section  through  a  Small  Artery  and  Vein  of  the  Mucous  Mem- 
brane of  a  Child's  Epiglottis;  the  artery  is  thick-walled  and  the  vein  thin-walled.  A, 
Artery;  the  letter  is  placed  in  the  lumen  of  the  vessel,  e,  Endothelial  cells  with  nuclei 
clearly  visible;  these  cells  appear  very  thick  from  the  contracted  state  of  the  vessel.  Outside 
it  a  double  wavy  line  marks  the  elastic  tunica  intima.  m,  Tunica  media  consisting  of 
unstriped  muscular  fibers  circularly  arranged;  their  nuclei  are  well  seen,  a,  Part  of  the 
tunica  advent'tia,  showing  bundles  of  connective-tissue  fiber  in  section,  with  the  circular 
nuclei  of  the  connective-tissue  corpuscles.  This  coat  gradually  merges  into  the  surrounding 
connective  tissue.  V,  The  lumen  of  the  vein.  The  other  letters  indicate  the  same  as  in 
the  artery.  The  muscular  coat  of  the  vein,  m,  is  seen  to  be  much  thinner  than  that  of  the 
artery,  "x  350.  (Klein  and  Noble  Smith.) 


are  gathered  up  into  larger  and  larger  trunks  until  they  terminate  in  the  two 
venae  cavse  and  the  coronary  vein  which  enter  the  right  auricle,  and  in  four 
pulmonary  veins  which  enter  the  left  auricle.  The  total  capacity  of  the 
veins  diminishes  as  they  approach  the  heart;  but  their  capacity  exceeds  by 
two  or  three  times  that  of  their  corresponding  arteries.  The  pulmonary 
veins,  however,  are  an  exception  to  this  rule.  The  veins  are  found  after 


178  THE    CIRCULATION    OF    THE   BLOOD 

death  more  or  less  collapsed  and  often  contain  blood.  They  are  usually 
distributed  in  a  superficial  and  a  deep  set  which  anastomose  frequently 
in  their  course. 

The  coats  of  veins  bear  a  general  resemblance  to  those  of  arteries,  figure 
149.  Thus,  they  possess  outer,  middle,  and  inner  coats.  The  outer  coat  is 
of  areolar  tissue  like  that  of  the  arteries,  but  is  relatively  thicker.  In  some 
veins  it  contains  a  few  musclar  longitudinal  cells.  The  middle  coat 
is  considerably  thinner  than  that  of  the  arteries;  it  contains  circular  un- 
striped  muscular  fibers  mingled  with  a  large  proportion  of  yellow  elastic  and 
white  fibrous  connective  tissue.  In  the  large  veins  near  the  heart  the 
middle  coat  is  replaced  for  some  distance  from  the  heart  by  circularly 
arranged  striped  muscular  fibers  continuous  with  those  of  the  auricles. 
The  internal  coat  of  veins  consists  of  a  fenestrated  membrane  lined  by 
endothelium.  The  fenestrated  membrane  may  be  absent  in  the  smaller 
veins. 


FIG.  150. — A,  Vein  with  valves  open.     B,  vein  with  valves  closed;  stream  of  blood  passing 
off  by  lateral  channel.     (Dalton.) 

The  veins  are  supplied  with  valves  in  certain  regions  of  the  body,  espe- 
cially in  the  arms  and  legs.  The  general  construction  of  these  valves  is 
similar  to  that  of  the  semilunar  valves  of  the  aorta  and  pulmonary  artery 
already  described.  Their  free  margins  are  turned  in  the  direction  toward 
the  heart,  so  as  to  prevent  any  movement  of  blood  backward.  They  are 
commonly  placed  in  pairs,  at  various  distances  in  different  veins.  In  the 
smaller  veins  single  valves  are  often  met  with,  and  three  or  four  are  some- 
times placed  together  or  near  one  another  in  the  larger  veins,  such  as  in  the 
subclavians  at  their  junction  with  the  jugular  veins.  During  the  period 
of  their  inaction,  when  the  venous  blood  is  flowing  in  its  proper  direction, 


ACTION    OF    THE    AURICLES  179 

they  lie  by  the  sides  of  the  walls  of  the  veins;  but  when  in  action  they  come 
together  like  valves  of  the  arteries,  figure  150.  Their  situation  in  the 
superficial  veins  of  the  forearm  is  readily  discovered  by  pressing  along  its 
surface,  in  a  direction  opposite  to  the  venous  current,  i.e.,  from  the  elbow 
toward  the  wrist,  when  little  swellings,  figure  150,  B,  will  appear  in  the 
position  of  each  pair  of  valves. 

Lymphatic  spaces  are  present  in  the  coats  of  both  arteries  and  veins; 
but  in  the  tunica  adventitia  or  external  coat  of  the  large  vessels  they  form 
a  distinct  plexus  of  more  or  less  tubular  vessels.  In  smaller  vessels  they 
appear  as  sinuses  lined  by  endothelium.  Sometimes,  as  in  the  arteries 
of  the  omentum,  mesentery,  and  membranes  of  the  brain,  the  pulmonary, 
hepatic,  and  splenic  arteries,  the  spaces  are  continuous  with  vessels  which 
distinctly  ensheath  them,  perivascular  lymphatic  sheaths.  Lymph  channels 
are  said  to  be  present  also  in  the  tunica  media. 

THE  ACTION  OF  THE  HEART. 

The  heart's  action  in  propelling  the  blood  consists  in  the  successive 
alternate  contraction,  systole,  and  relaxation,  diastole,  of  the  muscular  walls 
of  the  auricles  and  the  ventricles.  This  activity  furnishes  the  power  which 
keeps  the  blood  moving  through  the  arteries,  capillaries,  and  veins.  The 
heart  in  its  activity  is  like  a  great  force  pump  in  that  it  injects  a  certain 
quantity  of  blood  at  each  contraction  into  the  great  arteries.  Owing  to 
the  interaction  between  this  heart-beat  and  the  peripheral  resistance  to  the 
flow  of  blood,  together  with  the  elasticity  of  the  vessels  themselves,  a  high 
pressure  in  the  arteries  is  maintained  all  the  time.  The  heart's  contrac- 
tions pumping  against  this  high  arterial  tension,  are  sufficient  to  maintain 
a  constant  flow  of  blood  through  the  capillaries,  and  therefore  through  the 
active  tissues. 

The  heart  beats  at  an  average  rate  of  about  72  times  per  minute  during 
life.  Each  successive  contraction  really  begins  in  the  great  veins,  the 
superior  vena  cava  and  extends  over  the  auricles  and  ventricles  in  regular 
sequence.  The  contraction  of  each  successive  part  is  called  its  systole 
and  the  relaxation  its  diastole.  The  diastole  covers  the  period  of  active 
relaxation  of  the  muscle  and  the  pause  before  beginning  its  next  con- 
traction. Each  muscular  chamber  of  the  heart  may,  therefore,  be  said 
to  have  its  own  systole  and  diastole.  The  whole  series  of  events  from  the 
beginning  of  one  contraction  until  the  corresponding  event  in  the  next 
contraction  is  described  as  the  cardiac  cycle. 

Action  of  the  Auricles. — The  description  of  the  action  of  the  heart 
may  be  commenced  at  that  period  in  each  cycle  in  which  the  whole  heart  is 
at  rest.  The  heart  is  then  in  a  passive  state.  The  auricles  are  gradually 


180  THE   CIRCULATION   OF   THE  BLOOD 

filled  with  the  blood  flowing  into  them  from  the  veins,  and  a  portion  of  this 
blood  passes  at  once  through  the  auricles  into  the  ventricles,  the  opening 
between  the  cavity  of  each  auricle  and  that  of  its  corresponding  ventricle 
being  free  during  the  pause  of  the  entire  heart.  The  auricles,  however, 
receive  more  blood  than  at  once  passes  through  them  to  the  ventricles. 
Near  the  end  of  the  pause  they  become  passively  distended.  At  this 
moment  a  contraction  wave  begins  at  the  bases  of  the  venae  cavae  and,  run- 
ning down  over  the  walls  about  the  mouths  of  the  veins,  passes  to  the 
muscular  walls.  The  contraction  of  the  auricles,  the  right  and  left 
contracting  at  the  same  time,  forces  the  blood  into  the  ventricles. 

The  contraction  of  the  muscular  walls  at  the  mouths  of  the  great  veins 
and  of  the  sinus  region  maintains  a  condition  of  constriction  during  the 
time  of  the  auricular  contraction.  This  hinders  the  reflux  of  blood  from 
the  auricles  into  the  veins  during  the  auricular  systole.  Any  slight  re- 
gurgitation  from  the  right  auricle  is  limited  by  the  valves  at  the  junction  of 
the  subclavian  and  internal  jugular  veins  beyond  which  the  blood  cannot 
move  backward,  and  by  the  coronary  vein  which  is  supplied  with  valve-like 
fold  at  its  mouth.  The  force  of  the  blood  propelled  by  the  auricle  into  the 
ventricle  at  each  auricular  systole  is  transmitted  in  all  directions,  but, 
being  insufficient  to  open  the  semilunar  valves,  it  is  expended  in  distend- 
ing the  walls  of  the  ventricle. 

Action  of  the  Ventricles. — The  dilatation  of  the  ventricles  which 
occurs  during  the  latter  part  of  the  diastole  of  the  auricles,  is  completed  by 
the  forcible  injection  of  the  contents  of  the  latter.  The  ventricles,  now 
distended  with  blood,  immediately  begin  to  contract.  The  tricuspid  and 
mitral  valves  are  closed  by  the  initial  reflux  of  blood,  or  possibly  by  the 
currents  of  blood  formed  by  the  sudden  injection  of  the  ventricles  by  the 
auricular  contractions.  The  ventricular  systole  follows  the  auricular 
systole  so  closely  that  it  seems  continuous  with  it.  As  a  result  of  the  ventri- 
cular systole,  sufficient  pressure  is  produced  on  its  contents  to  overcome  the 
pressure  against  the  semilunar  valves  of  the  aorta  and  the  pulmonary 
artery,  and  the  ventricles  are  then  emptied  completely.  After  the  whole 
of  the  blood  has  been  expelled  from  the  ventricles,  the  walls  remain  con- 
tracted for  a  brief  period. 

The  form  and  position  of  the  fleshy  columns  on  the  internal  walls  of  the 
ventricles  no  doubt  help  to  produce  the  obliteration  of  the  ventricular 
cavities  during  contraction.  The  completeness  of  the  closure  may  often 
be  observed  on  making  a  transverse  section  of  a  heart  shortly  after  death 
in  any  case  in  which  rigor  mortis  is  very  marked,  figure  136.  In  such  a  case 
only  a  central  fissure  may  be  discernible  to  the  eye  in  the  place  of  the 
cavity  of  each  ventricle.  The  arrangement  of  the  muscles  of  the  heart, 
as  described  on  page  171,  is  such  as  to  expend  the  whole  force  of  the 
contraction  in  diminishing  the  cavity  of  the  ventricle,  or,  in  other  words, 
in  expelling  the  contents  of  blood. 


ACTION    OF    THE   VALVES  l8l 

On  the  conclusion  of  the  systole  the  ventricular  diastole  begins.  The 
muscular  walls  relax  and,  by  virtue  of  their  elasticity,  a  slight  negative 
pressure  may  be  set  up.  This  negative  or  suctional  pressure  on  the  left 
side  of  the  heart  may  be  of  importance  in  helping  the  pulmonary  circula- 
tion. It  is  somewhat  inconstant  in  appearance,  but  has  been  found  to  be 
equal  to  as  much  as  20  mm.  of  mercury,  and  is  said  to  be  quite  independent 
of  the  aspiratory  power  of  the  thorax  itself,  which  will  be  described  in  a 
later  chapter.  The  ventricles  now  remain  in  a  state  of  relaxation  or  rest 
until  the  next  systole  begins. 

The  duration  of  the  ventricular  systole  and  the  diastole  has  been  variously 
estimated.  A  computation  of  the  time  of  these  two  phases,  for  man,  in 
figure  153,  reproduced  from  Hiirthle,  gives  for  the  systole  0.38  of  a  second 
and  for  the  diastole  0.4  of  a  second,  with  a  total  of  o.  78  of  a  second.  This 
is  equivalent  to  a  rate  of  77  per  minute.  Variation  in  the  time  of  the  systole 
and  the  diastole  of  the  ventricle  falls  chiefly  on  the  pause  of  the  diastole. 

The  ventricles  undergo  little  or  no  change  of  shape  in  the  unopened  chest. 
At  the  moment  in  the  systole  when  the  ventricles  begin  to  discharge  their 

Conus  arteriosus 


Left  posterior  cusp  of 
pulmonary  valve 


Left  posterior  cusp  of 
aortic  valve 


Right  coronary 
artery 

Anterior  cusp  of 
aortic  valve 

Right  posterior  cusp 


Right  (marginal) 
cuspoUncusp,d 

Posterior  (septal) 

Posterior  cusp  of         v  -\,v  ^  ^  \  M^M  CUSP  °^  tricuspid 

mitral  valve  ?  :'  ValvC 

Left  ventricle 

Right  ventncle 


FIG.  151. — The  Bases  of  the  Ventricles  of  the  Heart,  showing  the  auriculo-ventricular, 
aortic,  and  pulmonary  orifices  and  their  valves.     (Cunningham.) 

contents  into  the  aorta  and  pulmonary  arteries,  respectively,  there  is  a  sharp 
decrease  in  size  of  the  ventricles.  This  decrease  takes  place  in  all 
dimensions. 

Action  of  the  Valves. — The  Tricuspid  Valve. — During  the  diastole 
of  both  auricles  and  ventricles  blood  flows  directly  through  the  auricles  into 
the  ventricles,  the  auricles  during  this  period  acting  as  continuations  of  the 
large  veins  which  empty  into  them.  At  the  end  of  the  period  the  ventricle 


1 82  THE    CIRCULATION    OF    THE   BLOOD 

on  each  side  has  already  been  filled  and  distended  by  the  pressure  of  blood 
from  the  veins.  The  systole  of  the  auricle  completes  this  filling  and  slightly 
overdistends  the  ventricle.  When  the  force  of  the  auricular  contraction  is 
spent,  the  ventricular  walls  rebound  slightly  toward  their  former  position 
and  in  so  doing  exert  some  pressure  upon  the  ventricular  side  of  the  tricuspid 
valve  which  floats  the  cusps  upward  toward  the  auricle.  In  this  connection 
another  force  comes  into  play,  viz.,  vortex  or  back  currents  resulting  from 
the  flow  of  the  blood  into  the  ventricle  under  the  pressure  of  the  auricular 
systole.  These  currents  aid  in  floating  the  valve  cusps  into  apposition. 
Thus  the  venous  orifices  of  the  ventricles  are  closed  at  the  end  of  the  auricular 
systole;  i.e.,  the  end  of  the  ventricular  diastole.  The  ventricular  systole 
which  follows  simply  serves  to  place  the  valves  under  greater  tension  thus 
closing  them  still  more  firmly.  It  should  be  recollected  that  the  diminution 
in  the  breadth  of  the  base  of  the  heart  in  its  transverse  diameters  during 


FIG.  152.— The  Tricuspid  Valves  of  the  Ox,  Closed.     Vertical  section.     (Krehl.) 

the  ventricular  systole  is  especially  marked  in  the  neighborhood  of  the 
venous  orifices,  and  this  aids  in  rendering  the  tricuspid  valve  competent  to 
close  the  openings  by  greatly  diminishing  the  diameter.  The  cusps  of  the 
valve  meet  not  by  their  edges  only,  but  by  the  opposed  surfaces  of  their  thin 
outer  borders.  The  margins  of  the  valve  are  still  more  secured  in  apposition 
with  one  another  by  the  simultaneous  contraction  of  the  papillary  muscles, 
whose  tendinous  chords  have  a  special  mode  of  attachment  for  this  very 
object.  They  compensate  for  the  shortening  of  the  ventricular  walls  and 
thus  prevent  the  valve  cusps  from  being  everted  into  the  auricles,  an  event 
that  does  occur  in  certain  valvular  lesions. 


THE  SOUNDS  OF  THE  HEART  183 

The  actions  of  the  tricuspid  and  mitral  valves  on  the  right  and  left  sides 
of  the  heart  are  essentially  the  same. 

The  Semilunar  Valves. — The  commencement  of  the  ventricular  systole 
precedes  the  opening  of  the  semilunar  valve  by  a  fraction  of  a  second.  The 
intraventricular  pressure  increases  with  the  progress  of  the  systole  until 
there  is  a  distinct  increase  over  the  arterial  pressure,  then  the  opening  of 
the  valves  takes  place  at  once.  The  valves  remain  open  as  long  as  this 
difference  continues.  When  the  diastole  of  the  ventricles  begins  and  the 
arterial  blood  pressure  exceeds  the  intraventricular  pressure,  there  is  an 
initial  reflux  of  blood  toward  the  heart  which  closes  the  semilunar  valve. 

The  dilatation  of  the  arteries  is  peculiarly  adapted  to  bring  this  about. 
The  lower  borders  of  the  semilunar  valves  are  attached  to  the  inner  surface 
of  the  tendinous  ring  which  bounds  the  orifice  of  the  artery.  The  tissue  of 
this  ring  is  tough  and  inelastic  and  the  valves  are  equally  inextensible, 
being  formed  mainly  of  tough  fibrous  tissue  with  strong  interwoven  cords, 
the  effect,  therefore,  of  each  propulsion  of  blood  from  the  ventricle  into  the 
artery  is  to  dilate  the  wall  of  the  first  portion  of  the  artery  and  the  three 
pouches  behind  the  valve  cusps  while  the  free  margins  of  the  cusps  are 
drawn  inward  toward  the  center.  This  position  of  the  valves  and  arterial 
walls  is  maintained  while  the  ventricle  continues  in  contraction;  but  as  it 
relaxes,  and  the  dilated  arterial  walls  recoil  by  their  elasticity,  the  blood  is 
forced  backward  toward  the  ventricles  and  onward  in  the  course  of  the 
circulation.  Part  of  the  blood  thus  forced  back  lies  in  the  pouches  (sinuses 
of  Valsalva)  between  the  valve  cusps  and  the  arterial  walls;  and  the  cusps 
are  pressed  together  till  their  thin  lunated  margins  meet  in  three  lines 
radiating  from  the  center  to  the  circumference  of  the  artery,  figure  151. 
The  corpora  Arantii  at  the  middle  of  the  free  margins  insure  a  more  effec- 
tive closure. 

The  Sounds  of  the  Heart. — When  the  ear  is  placed  on  the  chest  over 
the  heart,  two  sounds  may  be  heard  at  every  beat.  They  follow  in  quick 
succession  and  are  succeeded  by  a  pause  or  period  of  silence.  The  first 
sound  is  dull  and  prolonged;  its  commencement  coincides  with  the  impulse 
of  the  heart  against  the  chest  wall,  and  just  precedes  the  pulse  at  the  wrist. 
The  second  sound  is  shorter  and  sharper,  with  a  somewhat  flapping  char- 
acter. The  periods  of  time  occupied,  respectively,  by  the  two  sounds 
taken  together  and  by  the  pause  between  the  second  and  the  first  are 
unequal.  According  to  Foster,  the  interval  of  time  between  the  beginning 
of  the  first  sound  and  the  second  sound  is  0.3  of  a  second,  while  between 
the  second  and  the  succeeding  first  it  is  nearly  0.5  of  a  second,  see  figures 
153,  154,  and  167.  The  relative  length  of  time  occupied  by  each  sound,  as 
compared  with  the  other,  may  be  best  appreciated  by  considering  the 
different  forces  concerned  in  the  production  of  the  two  sounds.  In  one 
case  there  is  a  strong,  comparatively  slow  contraction  of  a  large  mass  of 


1 84 


THE    CIRCULATION    OF   THE   BLOOD 


muscular  fibers,  urging  forward  a  certain  quantity  of  fluid  against  con- 
siderable resistance;  while  in  the  other  it  is  a  strong  but  shorter  and  sharper 
recoil  of  the  elastic  coat  of  the  large  arteries — shorter  because  there  is  no 
resistance  to  the  flapping  back  of  the  semilunar  cusps  as  there  was  to  their 
opening.  The  sounds  may  be  expressed  by  the  words  lubb — dub.  The 
beginning  of  the  first  sound  corresponds  in  time  with  the  three  coincident 
events,  namely,  the  beginning  of  the  contraction  of  the  ventricles,  the 
closure  of  the  tricuspid  and  mitral  valves,  and  the  first  part  of  the  dilatation 
of  the  auricles.  The  sound  continues  through  a  somewhat  longer  interval 
than  the  second  sound.  The  second  sound,  in  point  of  time,  immediately 


FIG.  153. — Simultaneous  Tracings  of  the  Cardiac  Impact,  or  Cardiogram  (lower),  and 
the  Heart  Tones  (upper),  of  Man.  The  cross  strokes  at  the  beginning  of  the  cardiac  sound 
tracing  and  on  the  cardiogram  mark  the  synchronous  events.  (Hiirthle.) 

follows  the  cessation  of  the  ventricular  contraction,  and  corresponds  with 
the  commencing  dilatation  of  the  ventricles  and  the  opening  of  the  semi- 
lunar  and  mitral  valves,  figure  154. 

The  exact  cause  of  the  first  sound  of  the  heart  is  not  absolutely  known. 
Two  factors  probably  enter  into  it.  First,  the  vibration  of  the  semilunar  and 
mitral  valves  and  of  the  chordae  tendineae.  Second,  the  vibration  of  the 


FIG.  154. — Simultaneous  Tracings  of  the  Heart  Tone  and  Pulse  of  the  Carotid  in  the 
Dog.  Ai  and  A 2,  First  and  second  sounds;  P,  pulse;  S,  time  in  tenths  and  fiftieths  of  a 
second.  (Einthoven  and  Geluk.) 

muscular  mass  of  the  ventricles  themselves.     The  same  mechanical  condi- 
tions produce  equal  tension  on  the  ventricular  muscle  itself  and,  according 


THE    CARDIAC    IMPULSE  185 

to  the  second  view  this  is  sufficient  to  account  for  the  first  sound.  Looking 
upon  the  contraction  of  the  heart  as  a  simple  contraction  and  not  as  a  series 
of  contractions,  or  tetanus,  it  is  at  first  sight  difficult  to  see  why  there 
should  be  any  muscular  sound  when  the  heart  contracts. 

The  cause  of  the  second  sound  is  more  simple  and  definite  than  that  of 
the  first.  It  is  entirely  due  to  the  vibration  consequent  on  the  sudden  closure 
of  the  semilunar  valves  when  they  are  pressed  down  across  the  orifices  of 
the  aorta  and  pulmonary  artery.  The  influence  of  these  valves  in  producing 
the  sound  was  first  demonstrated  by  Hope  who  experimented  with  the  hearts 
of  calves.  In  these  experiments  two  delicate  curved  needles  were  inserted, 
one  into  the  aorta  and  another  into  the  pulmonary  artery  below  the  line  of 
attachment  of  the  semilunar  valves.  After  being  carried  upward  about 
half  an  inch  the  needles  were  brought  out  again  through  the  coats  of  the 
respective  vessels,  so  that  in  each  vessel  one  valve  was  held  back  against 
the  arterial  walls.  Upon  applying  the  stethoscope  to  the  vessels  it  was 
found  that  after  such  an  operation  the  second  sound  had  ceased  to  be  audible. 

Tube  to  communicate 
with  the  tambour 


Tympanum 


Ivory    Tape  to  attach 

knob        instrument  to  the  chest 


FIG.  155. — Cardiograph.     (Sanderson's.) 

Disease  of  these  valves,  when  sufficient  to  interfere  with  their  efficient  action, 
also  demonstrates  the  same  fact  by  modifying  the  second  sound  or  destroying 
its  distinctness. 

The  Cardiac  Impulse. — The  heart  may  be  felt  to  beat  with  a  slight 
shock  or  impulse  against  the  walls  of  the  chest  at  the  level  of  the  fifth  inter- 
costal space  on  the  left  side.  Its  extent  and  character  vary  in  different 
individuals,  a  factor  of  considerable  clinical  significance,  and  therefore  es- 
pecially discussed  in  works  on  clinical  diagnosis.  The  cause  of  the  cardiac 
impulse  has  been  variously  described.  It  will  be  remembered  that  during 
the  period  which  precedes  the  ventricular  systole  the  relaxed  heart  rests 
quietly  in  the  pericardial  cavity  and  with  its  apex  exerting  no  pressure 


1 86 


THE    CIRCULATION    OF    THE   BLOOD 


against  the  wall  of  the  chest.  When  the  ventricles  contract,  their  walls 
suddenly  become  firm  and  tense.  Being  firmly  attached  to  the  base  the 
effect  of  the  movement  is  to  press  the  hardened  ventricle  against  the 

Screw  to  adjust  the  lever 


Writing  lever  Tambour  Tube  to  the  cardiograph 

FIG.  156. — Marey's  Tambour,  to  which  the  Movement  of  the  Column  of  Air  in  the 
Cardiograph  is  Conducted  by  a  Tube,  and  from  which  it  is  Communicated  by  the  Lever 
to  a  Revolving  Cylinder  so  that  the  tracing  of  the  movement  of  the  cardiac  impulse  is 
obtained. 

chest  wall.  The  discharge  of  the  contents  of  the  ventricle  into  the  curved 
aorta  intensifies  this  pressure  by  its  mechanical  effect  in  tending  to 
straighten  the  curve  of  that  vessel  and  thus  holds  the  ventricle  in  firm 
contact  with  the  chest.  It  is  this  sudden  pressure  of  the  contracting 
heart  against  the  chest  wall  that  is  felt  on  the  outside.  The  impact  or 


FIG.  157. — Typical  Cardiogram  (upper  trace)  from  the  Dog.  Taken  simultaneously 
with  the  aortic  pressure  (middle)  and  intraventricular  pressure  (lower)  tracings.  Time 
in  o.oi  of  a  second.  (Hiirthle.) 

shock  is  possibly  more  distinct  because  of  the  partial  rotation  of  the 
whole  heart  toward  the  right  and  front  along  its  long  axis.  The  move- 
ment of  the  chest  wall  produced  by  the  ventricular  contraction  against 
it  may  be  registered  by  means  of  an  instrument  called  the  cardiograph; 


ENDOCARDIAC    PRESSURE 


I87 


and  the  record  or  tracing,  called  a  cardiogram,  corresponds  almost  ex- 
actly with  a  tracing  obtained  by  an  instrument  applied  over  the  con- 
tracting ventricle  itself. 

The  cardiograph,  figure  155,  consists  of  a  cup-shaped  metal  box  over 
the  open  front  of  which  is  stretched  an  elastic  india-rubber  membrane  upon 
which  is  fixed  a  small  knob  of  hard  wood  or  ivory.  This  knob,  however, 
may  be  attached,  as  in  the  figure,  to  the  side  of  the  box  by  means  of  a 
spring,  and  may  be  made  to  act  upon  a  metal  disc  attached  to  the  elastic 
membrane. 

The  knob  is  for  application  to  the  chest  wall  over  the  place  of  the 
greatest  impulse  of  the  heart.  The  box  or  tambour  communicates  by 
means  of  an  air-tight  tube  with  the 
interior  of  a  second  or  recording  tam- 
bour supplied  with-  a  long  and  light 
writing  lever  figure  156.  The  shock 
of  the  heart's  impulse  being  communi- 
cated to  the  ivory  knob,  and  through 
it  to  the  first  tambour,  the  effect  is, 
of  course,  at  once  transmitted  by  the 
column  of  air  in  the  elastic  tube  to 
the  interior  of  the  second  recording 
tambour,  also  closed,  and  through 
the  elastic  and  movable  disc  of  the 
latter  to  the  writing  lever  which  is 
adjusted  to  a  registering  apparatus. 
This  latter  generally  consists  of  a 
cylinder  or  drum  covered  with  smoked 
paper  and  revolves  by  clock-work 
with  a  definite  velocity.  The  point 
of  the  lever  writing  upon  the  paper 
produces  a  tracing  of  the  heart's  im- 
pulse, a  cardiogram. 

Endocardiac  Pressure. — The  effect 
of  the  muscular  contractions  and 
relaxations  of  the  walls  of  the  heart 
during  its  systole  and  diastole  is  to 
produce  changes  of  pressure  on  its 
content  of  blood.  When  this  pressure 
is  measured  by  the  proper  instrument 
it  is  found  that  the  pressure  in  the 
left  ventricle  varies  between  wide  ranges.  With  the  beginning  of  the 
muscular  contraction,  the  pressure  rises  till  it  slightly  exceeds  that  of  the 
pressure  of  the  aorta,  remains  high  for  a  brief  interval  of  time,  then  slowly 


FlG.  158.— Double  Cardiac  Sound 
for  Simultaneous  Registration  of  the 
Blood  Pressure  in  the  Right  Auricle 
and  Ventricle,  or  in  the  Aorta  and 
Left  Ventricle.  (Hurthle.) 


i88 


THE    CIRCULATION    OF    THE   BLOOD 


and  quietly  decreases  to  less  than  that  of  atmospheric  pressure.  It 
remains  low  until  the  beginning  of  the  next  systole.  For  the  right  ven- 
tricle the  events  and  variations  are  relatively  the  same. 


FIG.  159. — Simultaneous  Registration  of  Curves  of  the  Left  Intra ventricular  Pressure 
(lower),  the  Aortic  Pressure  (middle),  and  the  Cardiac  Impact  (upper).  Time,  o.oi  of  a 
second.  (Hiirthle.) 

A 


ft 


200 
ISO 


60 


0,1 


0,6 


0,7 


as       $eb 


Systole 


Diastole/. 


FIG.  1 60. — Schematic  Cardiogram  I,  and  Intraventricular  Pressure  Curves  II,  from  the 
Dog.  The  ventricular  pressure  curve  of  the  descending  type  is  represented  by  the  dotted 
line.  Pressure  in  millimeters  of  mercury,  time  in  tenths  of  a  second.  (Hiirthle.) 

In  order  to  determine  the  endocardiac  pressure  communication  must 
be  established  with  the  cavities  of  the  heart.  This  is  accomplished  by  a 
tube  known  as  a  sound,  which  is  introduced  into  the  left  ventricle  by 
passing  it  down  the  common  carotid  artery,  or  into  the  right  auricle  and 


ENDOCARDIAC    PRESSURE  1 89 

ventricle  through  the  jugular  vein.  When  a  cardiac  sound  is  introduced 
and  connected  with  some  form  of  pressure-recording  apparatus,  accurate 
tracings  of  the  variations  in  pressure  during  the  heart-beat  are  obtained. 
Chauveau  and  Marey  recorded  and  measured  with  accuracy  the 
variations  of  the  endocardiac  pressure  and  the  comparative  duration  of 


FIG.  161. — Apparatus  of  MM.  Chauveau  and  Marey  for  Estimating  the  Variations  of  Endo- 
cardiac Pressure,  and  Production  of  the  Impulse  of  the  Heart. 

the  contractions  of  the  auricles  and  ventricles.  They  placed  three  small 
india-rubber  air-bags  or  sounds  in  the  interior,  respectively,  of  the  right 
auricle,  the  right  ventricle,  and  in  an  intercostal  space  in  front  of  the  heart 
of  living  animals — the  horse.  These  bags  were  connected  by  means  of 


FIG.  162.— Tracings  of  i,  Intra-auricular;  2,  Intraventricular  Pressures;  and  3,  of 
the  Cardiac  Impulse  of  the  Heart.  To  be  read  from  left  to  right.  Obtained  by  Chauveau 
and  Marey. 

long  narrow  tubes  with  three  levers  arranged  one  over  the  other  in  con- 
nection with  a  registering  apparatus,  figure  161.  The  synchronism  of  the 
impulse  with  the  contraction  of  the  ventricles  is  also  well  shown  by 
means  of  the  same  apparatus,  and  the  causes  of  the  several  vibrations  of 
which  it  is  really  composed  have  been  demonstrated. 


THE    CIRCULATION    OF    THE   BLOOD 


In  the  tracing,  figure  162,  the  intervals  between  the  vertical  lines  rep- 
resent periods  of  a  tenth  of  a  second.  The  parts  on  which  any  given  vertical 
line  falls  represent  simultaneous  events.  It  will  be  seen  that  the  contraction 
of  the  auricle,  indicated  by  the  marked  curve  at  A  in  the  first  tracing,  causes 
a  slight  increase  of  pressure  in  the  ventricle  which  is  shown  at  A '  in  the  second 
tracing,  and  produces  also  a  slight  impulse,  which  is  indicated  by  A"  in  the 
third  tracing.  The  closure  of  the  semilunar  valves  causes  a  momentarily 
increased  pressure  in  the  ventricle  at  D',  affects  the  pressure  in  the  auricle  D, 
and  is  also  shown  in  the  tracing  of  the  cardiac  impulse  D". 

The  large  curve  of  the  ventricular  and  the  cardiac  impulse  tracings, 
between  A'  and  D',  and  A"  and  D",  are  caused  by  the  ventricular  contrac- 
tion, while  the  smaller  undulations,  between  B  and  C,  Bf  and  C',  B"  and  C", 
are  caused  by  the  vibrations  consequent  on  the  tightening  and  closure  of 
the  tricuspid  and  mitral  valves. 


FlG.  163. — Apparatus  for  Recording  the  Endocardiac  Pressure.      (Rolleston.) 

It  seems  by  no  means  certain  that  Marey's  curves  properly  represent 
the  variations  in  intraventricular  pressure.  Objection  has  been  taken  to 
his  method  of  investigation :  First,  because  his  tambour  arrangement  does 
not  admit  of  both  positive  and  negative  pressure  being  simultaneously  re- 
corded; second,  because  the  method  is  applicable  only  to  large  animals, 
such  as  a  horse;  third,  because  the  intraventricular  changes  of  pressure 


ENDOCARUIAC   PRESSURE 


IQI 


are  communicated  to  the  recording  tambour  by  a  long  elastic  column  of  air; 
and  fourth,  because  the  tambour  arrangement  has  a  tendency  to  record 
inertia  vibrations.     H.  D.  Rolleston,   who  has  pointed  out  the  above  in> 
perfections  of  Marey's  method,  has  reinvestigated  the  subject  with  a  more 
suitable  apparatus. 

The  method  adopted  by  Rolleston  is  as  follows: 

A  window  is  made  in  the  chest  of  an  anesthetized  and  curarized  animal,  and 
an  appropriately  curved  glass  cannula  introduced  through  an  opening  in  the 
auricular  appendix.  The  cannula  is  then  passed  through  the  auriculo-ventric- 


FIG.  164.— Endocardiac  Pressure  Curve  from  the  Left  Ventricle  of  the  Dog  The 
thorax  was  opened  and  a  cannula  introduced  through  the  apex  of  the  ventricle;  the  abscissa 
is  the  line  of  atmospheric  pressure.  G  to  D  represents  the  ventricular  contraction-  from 
D  to  the  next  rise  at  G  represents  the  ventricular  diastole.  The  notch,  at  the  top  of 'which 
is  F,  is  a  post-ventricular  rise  in  pressure  from  below  that  of  the  atmosphere  and  not  a 
presystohc  or  auricular  rise  in  pressure. 

ular  orifice  without  causing  any  appreciable  regurgitation  into  the  auricle, 
or  it  may  be  introduced  into  the  cavity  of  the  right  or  left  ventricle  by  an  opening 
made  in  the  apex  of  the  heart.  In  some  experiments  the  trocar  is  pushed  through 
the  chest  wall  into  the  ventricular  cavity.  His  apparatus  is  filled  with  a  solution 
of  leech  extract  in  0.75  per  cent,  saline  solution,  or  with  a  solution  of  sodium 
bicarbonate  of  specific  gravity  1.083. 


FIG.  165.— Curve  with  a  Dicrotic  Summit  from  the  Left  Ventricle;  the  Abscissa  Shows  the 

Atmospheric  Pressure. 

The  animals  employed  were  chiefly  dogs.  The  movement  of  the  column  of 
blood  is  communicated  to  the  writing  lever  by  means  of  a  vulcanite  piston  which 
moves  with  little  friction  in  a  brass  tube  connected  with  a  glass  cannula  by  means 
of  a  short  connecting  tube. 

When  the  lower  part  of  the  tube,  A,  is  placed  in  communication  with  one 


IQ2 


THE    CIRCULATION    OF    THE   BLOOD 


of  the  cavities  of  the  heart,  the  movements  of  the  piston  are  recorded  by  means 
of  the  lever,  C.  Attached  to  the  lever  is  a  section  of  a  pulley,  H,  the  axis  of  which 
coincides  with  that  of  the  steel  ribbon,  £;  while,  firmly  fixed  to  the  piston,  is 
the  curved  steel  piston  rod,  /,  from  the  top  of  which  a  strong  silk  thread,  J, 
passes  downward  into  the  groove  on  the  pulley. 

This  thread,  /,  after  being  twisted  several  times  around  a  small  pin  at  the 
side  of  the  lever,  enters  the  groove  in  the  pulley  from  above  downward,  and 
then  passes  to  be  fixed  to  the  lower  part  of  the  curve  on  the  piston  rod  as  shown  in 
the  smaller  figure. 

The  movement  of  the  lever,  C,  is  controlled  by  the  resistance  to  torsion  of 
the  steel  ribbon,  E,  to  the  middle  of  which  one  end  of  the  lever  is  securely  fixed 
by  a  light  screw  clamp,  F.  At  some  distance  from  this  clamp,  the  distance 
varying  with  the  degree  of  resistance  which  it  is  desired  to  give  to  the  move- 
ments of  the  lever,  are  two  holders,  Gt  Gr,  which  securely  clamp  the  steel 
ribbon. 

As  the  torsion  of  a  steel  wire  or  strip  follows  Hooke's  law,  the  torsion  being 
proportional  to  the  twisting  force,  the  movements  of  the  lever  point  are  pro- 
portional to  the  force  employed  to  twist  the  steel  strip  of  ribbon — in  other 
words,  to  the  pressures  which  act  on  the  piston,  B.  To  make  it  possible  to 
record  satisfactorily  the  very  varying  ventricular  and  auricular  pressures,  the 
resistance  to  torsion  of  a  steel  ribbon  adapts  itself  very  conveniently. 

This  resistance  can  be  varied  in  two  ways,  ist,  by  using  one  or  more  pieces 
of  steel  ribbon  or  by  using  strips  of  different  thicknesses;  or  ad,  by  varying 
the  distance  between  the  holders,  G,  G',  and  the  central  part  of  the  steel 
ribbon  to  which  the  lever  is  attached. 


FIG.  166. — Hiirthle's  Spring  Manometer.     A ,  Viewed  from  the  side;  B,  viewed  from  the  top. 

Rolleston's  conclusions  are:  That  there  is  no  distinct  and  separate 
auricular  contraction  marked  in  the  pressure  curves  obtained  from  the  right 
or  the  left  ventricle,  the  auricular  and  ventricular  rises  of  pressure  being 
merged  into  one  continuous  rise.  He  concludes  that  the  tricuspid  and 
mitral  valves  are  closed  before  there  is  any  great  rise  of  pressure  within  the  ven- 
tricle above  that  which  results  from  the  auricular  systole,  a,  figure  165.  The 


ENDOCARDIAC    PRESSURE 


193 


closure  of  the  valve  occurs  probably  in  the  lower  third  of  the  rise  AB,  figure 
165,  and  does  not  produce  any  notch  or  wave.  It  is  shown  that  the  semilunar 
valve  opens  at  the  point  in  the  ventricular  systole,  situated  at  c,  about  or  a 
little  above  the  junction  of  the  middle  and  upper  thirds  of  the  ascending  line 
AB,  and  the  closure  about  or  a  little  before  the  shoulder,  D,  The  figures 
show,  finally,  that  the  minimum  pressure  in  the  ventricle  may  fall  below 
that  of  the  atmosphere,  but  that  the  amount  varies  considerably. 

On  the  whole,  the  most  satisfactory  recording  instrument  for  the  measure- 
ment of  endocardiac  pressures  is  the  membrane  manometer  devised  by 
Hurthle.  This  instrument  avoids  mechanical  errors  in  a  most  satisfactory 
manner.  By  simultaneous  tracings  of  the  pressure  in  the  ventricle  and  in 
the  aorta  by  Hiirthle's  differential  manometer,  the  exact  moment  of  the 


PAHSE 

DIASTOLE 

AURICLE 

VENTRICLE 


IMPULSE 


FIG.  167. — Diagrammatic  Representation  of  the  Events  of  the  Cardiac  Cycle.  For 
events  which  occur  in  sequence,  read  in  the  direction  of  the  curved  arrow;  for  synchronous 
events,  read  from  the  center  to  the  periphery  in  any  direction.  (Coleman.) 

opening  and  closing  of  the  semilunar  valve  has  been  determined.  By 
similar  methods  we  have  been  able  to  fix  synchronism  between  other  events 
occurring  during  the  beat.  These  we  will  summarize  in  the  following  section. 


IQ4  THE    CIRCULATION    OF    THE   BLOOD 

Cardiac  Cycle. — The  entire  series  of  occurrences  in  a  single  heart- beat 
is  called  the  cardiac  cycle.  If  the  condition  of  the  heart  is  considered  at  that 
moment  when  its  muscular  walls  are  at  rest  it  will  be  found  that  the  tricuspid 
and  mitral  valves  are  open,  that  the  blood  is  flowing  from  the  great  veins 
into  the  auricle  and  the  ventricle,  which  form  a  continuous  cavity,  and 
that  the  pressure  is  about  that  of  the  atmosphere,  but  slowly  rising.  Now 
a  wave  of  contraction  begins  at  the  sinus  node  and  extends  over  the  auri- 
cles, which  immediately  contract  and  discharge  their  blood  into  the  ventri- 
cles, somewhat  distending  their  walls.  At  this  moment  the  ventricular 
systole  begins,  the  tricuspid  and  mitral  valves  are  closed,  the  flow  of  blood 
into  the  ventricles  is  checked,  and  the  first  heart  sound  is  heard.  The  con- 
traction of  the  ventricles  produces  a  rapidly  rising  pressure  on  the  enclosed 
contents  until  the  pressure  exceeds  that  in  the  pulmonary  artery  (and 
aorta),  the  semilunar  valves  open,  and  the  blood  is  discharged  into  the 
arteries.  The  ventricles  ordinarily  remain  contracted  for  a  brief  moment 
after  their  contents  are  emptied. 

The  ventricular  diastole  begins  next.  With  the  initial  relaxation  and 
the  first  slight  fall  of  the  intraventricular  pressure  below  that  of  the  aorta, 
the  semilunar  valves  close  and  the  second  sound  is  heard.  The  relaxation 
rapidly  proceeds  and  the  intraventricular  pressure  drops  to  below  atmos- 
pheric pressure,  the  auriculo- ventricular  valves  fall  open,  the  blood  that 
has  been  accumulating  in  the  auricles  flows  into  the  ventricles  and  the 
whole  heart  is  in  the  state  of  pause  described  as  the  point  of  beginning  of 
the  cycle. 

The  duration  of  the  cardiac  cycle  varies  with  the  heart  rate.  With  a 
rate  of  75  per  minute,  the  cardiac  cycle  will  take  o .  8  of  a  second.  In  round 
numbers  the  systole  of  the  auricle  takes  o .  i  of  a  second  with  a  diastole  of 
0.7  of  a  second,  o .  6  of  which  is  in  the  pause  or  rest  period.  The  ventricle 
requires  about  0.3  of  a  second  for  the  systole,  0.5  of  a  second  for  the  dias- 
tole, with  o .  2  to  o .  3  of  this  for  the  pause.  It  is  evident  that  the  whole  heart 
is  at  rest  at  the  same  instant  for  from  o .  i  to  o .  2  of  a  second. 

The  relations  of  the  cardiac  sounds  to  the  systole  and  the  diastole  have 
been  graphically  recorded  by  Hiirthle,  figure  153,  and  by  Einthoven  and 
Geluk,  figure  154.  The  former  found  that  in  a  heart-beat  lasting  o.  76  of  a 
second  the  interval  of  time  between  the  beginning  of  the  first  and  second 
sounds  was  0.25  of  a  second,  and  that  the  sounds  occur  just  at  the  begin- 
ning of  the  ventricular  systole  and  diastole,  respectively. 

During  the  cardiac  cycle  the  ventricles  are  completely  closed  from  the 
moment  of  the  beginning  of  the  ventricular  systole  until  the  pressure  amounts 
to  a  little  greater  than  the  pressure  in  the  corresponding  arteries,  which 
takes  about  0.2  of  a  second.  From  the  opening  of  the  semilunar  valves 
until  the  closure  of  those  valves,  about  o .  15  of  a  second,  the  ventricular  cavity 
is  in  open  communication  with  the  arteries.  There  is,  during  the  diastole, 


THE  PROPERTIES  OF  THE  HEART  MUSCLE 


195 


a  second  moment  of  complete  closure  of  the  ventricles,  from  the  time  of  the 
closing  of  the  semilunar  valves  until  the  ventricular  pressure  falls  below  the 
auricular  pressure  which  permits  the  tricuspid  and  mitral  valves  to  open. 

The  Force  of  the  Cardiac  Action. — In  estimating  the  amount  of 
work  done  by  a  machine  it  is  usual  to  express  it  in  terms  of  work  units.  A 
convenient  work  unit  for  this  purpose  is  the  amount  of  energy  required  to 
lift  a  unit  of  weight,  i.e.,  i  gram  or  i  kilogram,  through  a  unit  of  height;  i.e., 
i  centimeter  or  i  meter,  the  work  required  being  i  gramcentimeter  for 
small  units,  and  i  kilogrammeter  for  large  units,  respectively.  The 
average  work  done  by  the  heart  at  each  contraction  can  be  readily  com- 
puted by  multiplying  the  weight  of  blood  expelled  by  the  ventricles  by  the 
height  through  which  it  would  have  to  be  lifted  to  overcome  the  resistance 
to  its  discharge  from  the  cavities  into  the  arteries. 

The  quantity  of  blood  expelled  and  the  pressure  of  the  arteries  can  only 
be  estimated  for  man.  But  the  computations  from  indirect  observations 
on  other  mammals  indicate  that  the  quantity  of  blood  discharged  from 
each  ventricle  at  a  single  contraction  is  from  80  to  100  c.c.  The  pressure 
of  the  aorta,  see  page  221,  is  an  average  of,  say,  120  mm.  of  mercury,  or 
126  cm.  of  blood.  The  pressure  in  the  pulmonary  artery  is  much  less,  say, 
30  mm.  (20  to  40),  of  mercury  or  40  cm.  of  blood.  Collecting  these  facts, 
we  have  the  following  computation: 


Blood 
discharged. 

Against 
pressure 
column  of 
blood. 

Work  in 
gramcenti- 
meters. 

The  left  ventricle 

no    C.C. 

156  cm. 

i4,o4o 

The  right  ventricle       

oo    c.c. 

40  cm. 

3,600 

Total  .  . 

00     C.C. 

240  cm. 

i7,64o 

This  computation  shows  that  each  heart-beat  expends  17,640  gramcenti- 
meters  (17.64  grammeters)  of  work.  The  amount  of  energy  developed  in 
the  contractions  of  the  auricles  may  be  ignored  in  this  calculation,  which  is 
at  best  only  of  relative  value.  Calculations  based  on  the  determinations  of 
Vierordt,  also  other  earlier  determinations,  give  much  higher  figures  than 
are  presented  here. 

The  Properties  of  the  Heart  Muscle. — It  is  evident  that  if  we  are 
to  arrive  at  any  adequate  explanation  of  the  action  of  the  heart,  one  of  the 
first  questions  that  must  be  considered  is,  what  are  the  fundamental 
properties  of  heart  muscle  as  such? 


THE    CIRCULATION    OF   THE   BLOOD 


It  has  already  been  shown,  page  62,  that  the  muscular  fibers  of  the 
heart  differ  in  structure  from  skeletal  muscle  fibers  on  the  one  hand,  and 
from  unstriped  muscle  on  the  other,  occupying  an  intermediate  position 


•A.& 


FlG-  '68.  FlG>  I69. 

FIG.  168.— The  Heart  of  a  Frog  (Rana  esculenta),  from  the  Front.  V,  Ventricle;  Ad 
?E  k  *?  '  auricle;  B,  bulbus  arteriosus,  dividing  into  right  and  left  aorte 

FIG.  169.— The  Heart  of  a  Frog  (Rana  esculenta),  from  the  Back.  s.  v.,  Sinus  venosus 
opened;  c.  s.  s.,  left  vena  cava  superior;  c.  s.  d.,  right  vena  cava  superior;  c.  i.,  vena  cava 
mfenor;  y.  p.,  vena  pulmonales;  A.  d,,  right  auricle;  A.  s.  left  auricle;  A.  p.,  opening  of 
communication  between  the  right  auricle  and  the  sinus  venosus.  X  2^-3.  (Ecker.) 

between  the  two  varieties.  The  heart  muscle,  however,  possesses  a  prop- 
erty which  is  not  possessed  by  skeletal  muscle,  or  by  unstriped  muscle 
to  such  a  degree,  namely,  the  property  of  contracting  rhythmically. 

Rhythmicity. — The  property  of  rhythmic  contraction  is  shown  by  the 
action  of  the  heart  within  the  body;  its  systole  is  followed  by  its  diastole  in 


FIG.  170. — Automatic  Contractions  of  Sinus  Muscle  from  the  Terrapin's  Heart  in  0.7 
per  cent.  Sodium  Chloride.     Time  in  minutes.     (New  figure  by  L.  Frazier.) 

regular  sequence  throughout  the  life  of  the  individual.  The  force  and  fre- 
quency of  the  systole  may  vary  from  time  to  time  as  occasion  requires,  but 
there  is  no  interruption  to  the  action  of  the  normal  heart  or  any  interfer- 


THE  PROPERTIES  OF  THE  HEART  MUSCLE  IQ7 

ence  with  its  rhythmical  contractions.  Further,  in  an  animal  rapidly  bled 
to  death,  the  heart  continues  to  beat  for  a  time,  varying  in  duration  with 
the  kind  of  animal  experimentally  dealt  with 

* 

and  depending  on  whether  or  not  there  is  entire 

absence    of  blood    within  the  heart  chambers. 

Furthermore,  if  the  heart  of  an  animal  be  removed  ,  .S 

from  the  body,  it  still  continue  its  alternate  sys-  £ 

tolic  and  diastolic  movements  for  a  varying  time. 

Thus  we  see  the  power  of  rhythmic  contraction 

depends  neither  upon  connection  with  the  central 

nervous    system  nor  yet  upon  the  stimulation 

produced    by  the  presence  of  blood  within  its 

chambers.     Whether  or  not  rhythmicity  is  a  prop- 

erty  of  heart  muscle,  as  such,  was  conclusively 

settled  by  Gaskell  and  by  numerous  later  investi-  ~ 


gators  by  a  very  simple  experimental  procedure. 

Gaskell  cut  thin  strips  of  the  apex  of  the  ventricle     ^^^^HH|     £  £ 
of  the  terrapin,  which  is  free  from  the  nerve  cells, 
at  least  nerve  ganglia,  and  found  that  they  con-     ^1 
tracted  rhythmically  for  hours.     This  experiment 

has  become  a  classic  one  for  the  study  of  the  .2  £ 

cardiac  muscular  tissue.     Strips  of  cardiac  mus- 
cle cut  from  the  auricle  and  from  the  contractile  H 
walls  of  the  venae  cavae,  or  sinus  venosus,  of  the                             '      •£  £ 
terrapin  also  contract  rhythmically.    If  the  strips 
of  muscle  are  kept  moist  with  the  same  blood     ^^^^_ , 
or  serum,  then  the  rhythm  of  the  sinus  is  greater                                   j~  if  I 
than  that  of  the  auricle,  and  that  of  the  auricle  g  ;| ' 
greater  than  that  of  the  ventricle,  a  difference                                   ,§  £.S 
that  is  based  on  a  physiological  differentiation 
of   the  tissue.     The  sinus  muscle  is  also  more     _^_^^^^^^_ 
delicately  responsive  to  stimuli  than  is  the  ven-                        M        *o  -0 
tricular  muscle;  i.e.,  it  is  more  irritable.                                      - 

Porter  first  performed  the  more  difficult  ex-     ^^^^^^^mm      «•  #-& 
periment  of  isolating  a  small  disc  of  muscle  from 

the  ventricle  of  the  dog,  leaving  only  the  delicate          -  .      ,        J "Sis 

nutrient  artery  through  which  the  muscle  was 
fed  with  defibrinated  blood.     This  isolated  small  \  j|  ,\ 

piece  of  ventricle  contracted  vigorously  for  many  t 

minutes.     Moorhouse  has  recently  shown  that 

various  portions  of  the  auricle,  the  sinus,  and  .;    .r  S  °  ^ 

the  veins  contract  in  good  rhythm.     They  also  ,,;--;          £  §•§ 

respond    to   various    drugs   in    a  characteristic     HHHEBHU 


198  THE    CIRCULATION    OF    THE   BLOOD 

fashion.     We  may  safely  conclude,  therefore,  that  the  mammalian  heart 
muscle  is  also  automatically  rhythmic. 

Tonicity. — Cardiac  muscle  is  characterized  by  its  maintaining  a  con- 
stant degree  of  partial  contraction  described  as  muscle  tone,  or  tonicity. 


FIG.  172.— Automatic  Contractions  of  a  Strip  of  Ventricular  Muscle  from  the  Apex 
of  the  Terrapin's  Heart  contracting  in  0.7  per  cent.  Sodium  Chloride;  from  +  to  +  0.03 
per  cent.  Potassium  Chloride  is  added  to  the  Sodium  Chloride.  The  thythm  is  recovered 
very  slowly  when  the  muscle  is  returned  to  0.7  per  cent,  sodium  chloride.  Time  in 
minutes  (upper)  and  seconds  (lower  stroke).  (Watkins  and  Elliott.) 

This  property  is  possessed  by  all  parts  of  the  heart.  In  the  auricle,  how- 
ever, and  especially  in  the  muscular  walls  of  the  sinus  and  veins,  there  is 
considerable  variation  in  tonicity.  Botazzi  showed  that  in  the  auricle  of 


FIG.  173. — Automatic  Contractions  of  a  Strip  of  Ventricular  Muscle  from  the  Apex 
of  the  Terrapin's  Heart,  a,  Contracting  in  0.7  per  cent,  sodium  chloride;  b,  when  0.03 
per  cent,  calcium  chloride  solution  is  added.  Time  in  minutes.  (Frazier.) 

the  toad  the  variations  of  tone  were  wave-like  and  periodic,  even  though 
the  auricle  were  contracting  with  its  ordinary  fundamental  rhythm. 
Howell  has  published  numerous  experiments  showing  tone  waves  in  auri- 
cular and  sinus  muscle  of  the  terrapin,  in  which  muscle  there  may  or  may 


CARDIAC    CONTRACTIONS  199 

not  be  occurring  at  the  same  time  the  ordinary  fundamental  rhythmic 
contractions,  figure  170. 

Irritability  of  Heart  Muscle. — Mention  was  made  above  of  the  difference 
in  irritability  of  heart  muscle  chosen  from  different  parts  of  the  heart. 
The  irritability  of  the  muscle  of  each  part  also  varies  during  the  different 
stages  of  the  contraction.  Experiment  shows  that  the  muscle  is  not  irri- 
table to  a  stimulus  applied  at  any  time  from  the  beginning  of  the  contrac- 
tion until  the  summit  of  the  contraction  is  reached.  This  is  called  the 
refractory  period.  From  the  summit,  through  the  relaxation  and  succeed- 
ing pause,  the  irritability  rapidly  increases  until  the  beginning  of  the 
next  contraction.  .  Considering  the  automatically  contracting  muscle,  the 
point  in  which  the  automatic  contraction  is  released,  i.e.,  begins,  is  the 
point  of  maximal  irritability.  It  is  the  moment  when  the  irritability  is 
so  great  that  the  muscular  equilibrium  is  no  longer  stable,  and  the  physio- 
logical contraction  results. 

The  irritability  of  heart  muscle  is  very  sharply  influenced  by  its  condition 
of  nutrition,  especially  by  the  inorganic  salts  present  in  the  blood  and  lymph, 
see  page  207.  The  salt  content  of  the  blood  comprises  about  0.7  per  cent, 
sodium  chloride,  0.03  per  cent,  potassium  chloride,  and  0.025  to  0.03  per 
cent,  calcium  (phosphate  probably),  as  well  as  traces  of  other  metal  bases. 


FIG.  174. — Refractory  Period  in  the  Ventricular  Strip  of  the  Terrapin. 

The  heart  muscle  has  been  shown  by  numerous  investigators  to  be  delicately 
responsive  to  the  proportions  of  these  salts  in  the  blood,  or  in  any  artificial 
solution  which  may  be  substituted  for  blood.  If  the  rhythm  is  to  be  taken 
as  an  index  of  the  irritability,  then  an  increase  of  sodium  and  calcium  salts 
increases  the  irritability  (rhythm),  while  the  influence  of  an  increase  in  potas- 
sium is  to  depress  the  irritability. 

Cardiac  Contractions  Always  Maximal. — The  heart  muscle  exhibits 
another  property  which  distinguishes  it  from  ordinary  skeletal  muscle,  viz., 
the  way  in  which  it  reacts  to  stimuli.  The  latter,  Chapter  XIII,  reacts 
slightly  to  a  stimulus  little  above  the  minimal,  and  with  an  increase  of  the 


200  THE   CIRCULATION    OF   THE  BLOOD 

strength  of  the  stimulus  will  give  contractions  of  increasing  amplitude  until 
the  maximum  contraction  is  reached.  In  the  case  of  the  heart-beats  this  is 
not  so,  since  the  minimal  stimulus  which  has  any  effect  is  followed  by  the  maxi- 
mum contraction;  in  other  words,  the  weakest  effectual  stimulus  brings  out  as 
great  a  contraction  as  the  strongest.  If  a  contraction  is  induced  earlier  than 
it  would  automatically  occur,  then  the  succeeding  pause  is  longer;  i.e.,  there 
is  a  compensatory  pause.  Also  the  contraction  induced  is  smaller  and  the 
one  following  the  compensatory  pause  is  proportionately  larger.  This 
observation  can  easily  be  demonstrated  on  the  heart  strip,  see  figure  174, 
or  on  the  whole  ventricle  of  the  frog,  which  was  originally  used  by 
Bowditch. 

Nerve  influence,  nutrition,  temperature,  etc.,  will  of  course  affect  the 
extent  of  the  contractions,  but  under  a  given  set  of  conditions  it  is  held  that 
the  contractions  which  occur  are  maximal  for  the  particular  set  of  nutri- 
tive and  other  conditions.  This  is  more  readily  understood  when  taken  in 
connection  with  the  fact  that  when  a  contraction  originates  in  a  cardiac 
cell  it  is  conducted  throughout  the  continuity  of  all  the  cells  of  the  mass. 

Theories  of  the  Heart-beat. — The  cause  of  the  rhythmic  power  of 
the  heart  as  a  whole  has  been  the  subject  of  much  discussion  and  experi- 
mental observation.  Two  leading  hypotheses  have  given  inspiration  to 
investigators,  and  now  one,  now  the  other  theory  has  attracted  followers 
as  new  facts  have  been  discovered.  These  are  known  as  the  neurogenic 
theory  and  the  myogenic  theory,  respectively,  though  neither  is  proven 
correct. 

The  heart  has  long  been  known  to  have  the  power  of  rhythmic  contrac- 
tions after  removal  from  the  body  and  after  all  connection  with  the  central 
nervous  system  has  been  destroyed. 

The  heart  can  be  taken  entirely  away  from  the  body  of  an  animal 
and  kept  beating  rhythmically  with  ease.  This  is  true  for  many  inverte- 
brates and  for  all  vertebrates  examined  including  fishes,  frogs,  turtles, 
snakes,  birds,  and  numerous  mammals  including  man  himself.  It  is 
only  necessary  to  supply  the  heart  through  its  local  circulatory  vessels 
with  the  proper  nutritive  fluid  well  aerated  with  oxygen  and  at  the 
normal  temperature  of  the  animal  from  which  the  heart  is  taken.  The 
question  long  debated  is  this.  What  initiates .  these  wonderfully  persis- 
tent and  regularly  repeated  contractions? 

If  the  frog's  or  terrapin's  heart  is  removed  from  the  body  entire,  it  will 
continue  to  contract  for  many  hours  and  even  days,  and  the  contractions 
have  no  apparent  difference  from  the  contractions  of  the  heart  before 
removal.  The  contractions  will  take  place,  as  we  have  mentioned,  without 
the  presence  of  blood  or  other  fluid  within  its  chambers.  Not  only  is  this 
the  case,  but  the  auricles  and  ventricle  may  be  cut  off  from  the  sinus, 
and  all  parts  continue  to  pulsate.  Further,  the  auricles  may  be  divided 


THEORIES    OF    THE    HEART-BEAT 


2OL 


from  the  ventricle  with  the  same  result.  If  the  heart  be  divided  lengthwise, 
its  parts  will  continue  to  pulsate  rhythmically.  The  ventricle  remains 
comparatively  quiet,  contractions  occurring  at  longer  intervals,  if  at  all. 
However,  the  isolated  ventricle  remains  irritable  so  long  as  bathed  in 
blood  or  in  a  balanced  Ringer  solution,  and  will  contract  upon  receiving 
a  slight  stimulus.  In  fact,  a  single  stimulus  will  often  call  forth  a  series  of 
contractions  of  the  ventricle.  The  frog's  ventricle,  when  its  muscular 


FIG.  175.— Isolated  Nerve  Cells  from  the  Frog's  Heart.     7,  Usual  form;  II,  twin  cell;  Ct 
capsule;  A7",  nucleus;  P,  process.     (From  Ecker.) 

and  nervous  connections  with  the  auricle  are  physiologically  severed,  as 
by  crushing,  will  remain  quiet  when  fed  by  its  own  blood,  though  it  will 
contract  rhythmically  when  fed  with  physiological  salt  solution. 

It  is  thus  seen  that  the  rhythmical  movements  of  these  parts  of  the 
heart  appear  to  be  more  marked  in  the  parts  at  the  venous  end  of  the 
organ,  i.e.,  the  sinus  and  auricle,  and  less  marked  in  the  ventricular  end. 
Ventricular  pieces  contract  when  still  connected  with  the  auricles  but  do 
not  readily  contract  in  the  ordinary  condition  even  when  irrigated  with 
blood.  These  are  regarded  as  facts  peculiarly  in  favor  of  the  view  that 
the  rhythm  is  inherent  in  the  special  nervous  elements  of  the  heart. 

This  view  which  has  long  been  known  as  the  neurogenic  theory,  attri- 
butes the  remarkable  power  of  the  heart  to  continue  contractions  after 
removal  from  the  body,  and  presumably  while  in  the  body,  to  the  presence 
of  the  collections  of  nerve  cells  within  the  walls  of  the  heart  itself.  The 
local  nervous  mechanism  in  the  frog  consists  of  three  chief  groups  of  cells 
or  ganglia.  The  first  group  is  situated  in  the  wall  of  the  sinus  venosus  at 
the  junction  of  the  sinus  with  the  right  auricle,  Remak's  ganglion.  The 
second  group  is  placed  near  the  junction  between  the  auricles  and  ven- 
tricles, Bidder's  ganglion.  The  third  is  in  the  septum  between  the  auricles, 
wn  Bezold's  ganglion.  Small  ganglia  have  been  described  for  the  base  of 
the  ventricle,  but  no  ganglia  are  present  in  the  apical  part  of  the  ventricles, 
though  isolated  neurones  have  been  found.  The  nerve  cells  of  which 
these  ganglia  are  composed  are  generally  unipolar,  seldom  bipolar.  Some- 


202  THE   CIRCULATION   OF   THE  BLOOD 

times  two  cells  are  said  to  exist  in  the  same  envelope,  constituting  the  twin 
cells  of  Dogiel.  The  cells  are  large,  and  have  very  large  round  nuclei  and 
nucleoli,  figure  175.  The  neurogenic  theory  assumes  that  the  periodic 
discharge  of  motor  nerve  impulses  takes  place  from  these  neurones  thus 
stimulating  the  musculature  of  the  heart  to  rhythmic  contractions.  The 
stimuli  start  at  the  region  of  the  sinus  and  are  conducted  over  the  heart 
in  orderly  sequence,  but  their  origin  in  nerves  is  questioned. 

In  the  myogenic  theory  ,the  heart's  rhythmical  contractions  are 
explained  as  due  to  the  inherent  property  of  the  cardiac  muscle  itself. 
Most  convincing  facts  in  support  of  this  theory  have  been  arrived  at  by  a 
study  of  cardiac  muscle,  as  such,  and  by  studies  on  the  whole  heart, 
particularly  by  Gaskell's  method  of  blocking.  The  term  blocking  is 
explained  as  follows:  It  will  be  remembered  that  under  normal  con- 
ditions the  wave  of  the  contractions  in  the  heart  starts  at  the  sinus  and 
travels  down  over  the  auricles  to  the  ventricles.  The  irritability  of  the 
muscle  and  its  power  of  rhythmic  contractions  is  greatest  in  the  sinus,  less 
in  the  auricles,  and  least  in  the  ventricles.  By  an  arrangement  of  liga- 
tures or  by  a  system  of  clamping,  one  part  of  the  heart  may  be  more  or  less 
isolated  from  any  other  portion.  With  such  a  clamp  the  contraction 
waves  can  be  more  or  less  completely  interrupted  in  their  passage  from  the 
sinus  end  of  the  heart  past  the  clamp  toward  the  ventricular  end.  If  the 
clamp  is  complete,  so  as  to  interrupt  the  physiological  continuity  between 
the  parts,  then  any  contractions  in  the  apical  portion  will  be  entirely 
independent  of  those  in  the  sino-auricular  portion.  If  the  blocking  is 
partial  only,  then  the  ventricular  end  of  the  heart  ordinarily  contracts 
in  unison  with  the  sino-auricular,  although  its  rate  may  be  as  i  to  2, 
i  to  3,  etc.  In  other  words,  only  every  second  or  every  third  sino-auricu- 
lar contraction  will  be  able  to  pass  the  block  to  the  ventricle. 

The  effects  of  blocking  are  due  to  the  interruption  of  muscle  continuity 
rather  than  to  nerve  continuity.  This  is  beautifully  demonstrated  by  an 
experiment  of  zigzag  cutting  of  the  ventricle  in  the  terrapin,  since  it  cannot 
be  supposed  that  any  nerves  would  pass  through  the  ventricular  mass  by 
such  a  zigzag  course.  In  this  experiment  the  contraction  wave  passes 
down  over  the  muscle  and  around  the  end  of  the  cuts  until  it  reaches  the 
apex.  The  apex  muscle  contracts  in  sequence  with  the  auricle  and 
successive  pieces  of  the  ventricle.  If  the  zigzag  cuts  are  made  almost 
complete  so  as  to  reduce  to  a  minimum  the  muscular  tissue  which  bridges 
from  one  cut  to  the  next,  then  it  happens  that  occasional  contractions  will 
be  unable  to  pass  and  the  apex  contracts  after  its  preceding  piece  in  the 
ratio  of  i  to  2,  or  i  to  3,  etc.,  as  described  above  Thus,  division  of 
the  muscle  has  the  same  effect  as  partial  clamping  in  the  same  position. 
These  facts  all  point  to  a  greater  power  of  rhythmicity  in  the  cardiac 
tissue  nearer  the  venous  end  of  the  heart.  This  difference  of  rhythmicity 


THEORIES    OF    THE  .HEART-BEAT  203 

is  not  due  to  the  nerves  of  the  heart,  say  the  myogenists,  but  to  the 
inherent  property  of  the  muscle  itself. 

It  was  thought  for  a  long  time  that  in  the  mammal  there  was  no  mus- 
cular continuity  between  the  auricles  and  ventricles  to  conduct  the  con- 
traction wave  and  that  this  was  an  insurmountable  difficulty  in  the  way  of 
accepting  the  myogenic  theory  of  the  heart  beat.  In  1893  Kent  described 
a  bundle  of  muscle  fibers  arising  in  the  wall  of  the  right  auricle  and  near  the 
septum  and  running  down  into  and  forming  a  muscular  connection  with  the 
ventricles.  This  bundle  was  also  independently  described  by  His,  Jr., 
and  generally  bears  his  name.  This  band  is  called  the  auriculo- ventricu- 
lar bundle. 

It  is  now  generally  recognized  that  the  early  embryonic  cardiac  tissue 
undergoes  differentiation  in  two  directions.  Out  of  one  of  these  types 
of  tissue  there  is  produced  the  wellknown  cardiac  muscular  tissue  which 
makes  up  the  mass  of  the  auricles  and  the  ventricles.  Out  of  the  other 
differentiation  is  produced  the  type  of  tissue  which  constitutes  the  auri- 
culo-ventricular  conducting  or  bundle  system.  This  conducting  type 
of  tissue  is  striated  like  the  ordinary  cardiac  tissue  but  in  general  appear- 
ance is  more  embryonic  in  type.  Its  cells  constitute  what  is  known  as  the 
Purkinje  fibers.  The  main  bundle  described  by  His,  Jr.,  runs  in  the  inter- 
ventricular  septum  somewhat  lightly  buried  in  the  tissue  beginning  at  the 
base  of  the  auricle  on  the  right  side  and  running  down  through  the  inter- 
auricular  tissue  to  the  septum  of  the  ventricles  where  it  divides  into  a 
right  and  left  branch.  Strands  of  this  tissue  extend  somewhat  up  into 
the  auricles  but  are  elaborately  developed  into  a  net  work  lying  just  under- 
neath the  endothelium  of  the  right  and  left  ventricles.  The  branch- 
ing net  work  of  these  cells  shades  into  and  is  continuous  with  cells  of  the 
ordinary  cardiac  type.  Miss  De  Witt  (1909)  made  an  excellent  model 
of  this  system  which  has  become  classic  in  the  literature  and  is  reproduced 
in  figure  106.  The  bundle  system  contains  two  regions  known  as  nodes, 
the  sino-auricular  node  imbedded  in  the  wall  of  the  right  auricle  just  in 
the  angle  where  it  is  joined  by  the  superior  vena  cava;  and  the  auriculo- 
ventricular  node,  described  first  by  Tawara,  which  lies  in  the  upper  end  of 
the  His  bundle.  The  physiological  differentiation  of  this  tissue  is  in  the 
direction  of  rhythm  production  and  facilitated  conduction. 

The  demonstration  of  the  auriculoventricular  bundle  has  proven  to  be 
of  the  strongest  support  to  the  myogenic  theory.  Erlanger  has  shown, 
by  an  ingenious  device  for  partially  clamping  this  muscular  band,  that 
even  the  mammalian  ventricle  exhibits  the  phenomenon  of  heart  block. 
The  sequence  of  auricle  and  ventricle  can  be  perfectly  controlled  by  the 
degree  of  compression  exerted  by  the  clamp.  In  his  experiments  the 
ventricle  contracts  in  unison  with  every  auricular  contraction,  or  only 
every  second  or  every  third,  according  to  the  degree  of  blocking. 


204 


THE    CIRCULATION    OF    THE   BLOOD 


It  was  shown  along  ago  (by  Merunowicz  in  1875)  that  the  isolated  apex 
of  the  ventricle  of  the  frog  remains  quiet  when  filled  with  blood,  but  readily 
gives  good  rhythmic  contractions  in  physiological  saline  and  other  artificial 
solutions.  The  inactivity  in  blood  is  not  necessarily,  therefore,  due  to 
nervous  isolation  from  the  ganglionated  parts  of  the  heart  nor  to  the 
bundle  system  alone.  Contractions  occur  in  the  small  bits  of  ventricular 
muscle  as  isolated  by  Gaskell,  and  these  may  continue  for  hours.  It  is. 
well  known  also  that  the  embryonic  heart  contracts  rhythmically  before 
nerve  cells  have  reached  the  organ  or  even  before  any  blood  is  formed,  as 
shown  in  the  embryos  of  certain  fishes. 


FIG.  176. — Stereoscopic  photograph  of  a  model  of  the  atrioventricular  nodal  system, 
in  the  calf's  heart.  Viewed  from  behind.  The  auricular  network  is  not  shown.  Should 
be  examined  through  a  stereoscope.  (Lydia  M.  DeWitt.) 

The  phenomena  of  heart  block,  the  contractions  of  the  ventricular  apex 
when  physiologically  isolated  from  the  parts  of  the  heart  which  contain  the 
ganglia,  the  behavior  of  isolated  strips  of  the  heart,  especially  of  the  ven- 
tricle and  the  rhythm  of  the  embryonic  heart  are  all  held  to  be  in  favor  of 
the  myogenic  theory.  However,  in  light  of  recent  developments  we  must 
find  our  explanation  of  the  block  phenomena  as  well  as  of  other  facts  used 
in  argument  for  the  myogenic  theory  in  the  physiology  of  this 
differentiation  from  the  embryonic  muscle,  namely,  the  bundle  system. 

PRESENT    VIEWS   OF  THE   AUTOMATICITY   OF  THE  HEART. 


Whatever  view  one  adopts  of  the  heart's  beat  he  has  to  explain  not  only 
the  periodic  origin  of  the  rhythm  but  also  to  explain  the  orderly  sequence 
of  auricles  and  ventricles.  Keith  and  Flack  (1906)  have  ascribed  the 
initial  rhythm  to  a  center  or  node,  the  sino-auricular  node  as  given  above. 
In  the  mammalian  heart  the  normal  beat  under  normal  conditions  is 


THE  AUTOMATICITY  OF  THE  HEART 


205 


generated  at  this  point  and  conduction  proceeds  in  an  orderly  manner  in  all 
directions  not  only  toward  the  ventricle  but  out  over  the  atria  and  on  to  the 
veins  themselves.  These  last  points  have  been  most  carefully  studied  and 


FIG.  1760.  —  Normal  electrocardiogram  of  man,  lead  II. 

established  by  Lewis  and  a  number  of  his  associates.     Tawara,  Eyster  and 

Meek,  Wilson,  Greene  and  Gilbert,  and  others  have  explained  that  under 

certain  conditions  rhythm  may  arise  lower  down  in  the  conducting  tissue, 

namely,  at  the  auriculo-ventricular  node  or  center  or  even  as  low  as  the 

bundle  branch  (Greene  and  Gilbert).     In  this  case  the  conduction  is  from 

the  point  of  rhythm  production  not 

only   toward   the  ventricles  which 

contract  together  in  response  to  the 

stimulus   reaching   them   from  the 

auriculo-ventricular  node,  but  con- 

duction is  reversed  toward  the  auri- 

cle leading  to  delay  in  the  auricular 

contraction  in  comparsion  with  the 

normal. 

It  is  obvious  that  the  tissue  of 
the  bundle  system  is  differentiated 
highly  in  the  direction  of  rhythm 
production  and  of  conductivity. 
When  once  the  rhythm  arises  in  the 
sino-auricular  node  the  stimulus  is 
conducted  five  to  ten  times  more 
rapidly  over  the  bundle  system  to 


FIG.  1766.—  The  figure  gives  the  times 
the  nroner  auricular  tissue  and  to    of  begmning  contraction  of  the  respective 

> 
the    larger    ventricular    mass    than 


points  measured  in  fractions  of  a   second 
after  the  beginning  of  the  R  in  lead  II.     The 


mass 

would  be  possible  through  the  slower 

•conducting    cardiac    muscle.     This    area  and  occurs  latest  near  the  base  of  the 
insures  the  contraction  of  the  entire  aorta-    (Lewis-) 

ventricular  walls  at  more  nearly  the  same  instant  than  would  otherwise 
be  the  case.     In  fact,  the  arrangement  of  muscle  bands  of  the  ventricle 


206 


THE    CIRCULATION    OF    THE    BLOOD 


does  not  materially  influence  the  spread  of  the  conduction  wave  over 
the  ventricular  walls  (Lewis)  as  was  once  supposed.  The  Purkinje  system 
furnishes  the  shorter  pathway.  These  facts  have  been  demonstrated 


0350 


0250 


•0120 


0200 


0250 


FIG.  176^. — Transection  of  the  dog's  heart  to  show  the  spread  of  the  conduction 
stimulus  from  the  bundle  to  the  different  parts  of  the  ventricular  walls,  and  the  delay 
in  conduction  through  the  walls  by  muscle  paths.  (Lewis.) 

by  recording  the  time  of  arrival  of  the  stimulus  at  different  points 
on  the  ventricular  wall,  figure  176^.  The  companion  figure,  176^,  gives 
a  schematic  transection  of  the  dog's  heart  to  indicate  the  direction 


FIG.  177. — The  rate  of  different  isolated  strips  of  mammalian  ventricle  under 

the  influence  of  changes  of  temperature.     —  —  nodal  strip,    

coronary  strip,  J^  temperature  variation,  time  in  minutes.    Tempera- 

ture in  degrees  centigrade,  rate  in  beats  per  ten  seconds.     (Moorehouse.) 

of  spread  of  the  stimulus  from  the  inner  surface  into  and  through  the 
muscle  walls  of  the  two  ventricles.  The  delay  in  sequence  between  the 
auricles  and  ventricles  is  represented  by  this  rate  of  conduction  through 


£ 


THE  AUTOMATICITY  OF  THE  HEART  '  207 

the  bundle  system.  It  is  approximately  0.16  seconds.  The  time  at 
which  the  contraction  stimulus  arrives  at  different  points  on  the  ventri- 
cles, measured  from  the  moment  of  the  beginning;  of  the  R  wave  is  illus- 
trated in  Figure  176^. 

This  newer  conception  of  the  heart's  differ- 

entiations  gives  to  the  true  cardiac  muscle  a     HHE!  *i£      ~' 

position  in  energy  production  under  the  direct  |:|* 

control  of  the  conducting  system  for  its  coor-     ^M  1 

dination.  To  the  conducting  system  is  rele- 
gated  the  function  of  stimulus  production  and 

a  rapid  distribution  of  the  stimulus  that  still     1?£:  £ 

~  ~ 
preserves  a  mechanically  efficient  sequence. 

The   function    of   the   nerve   elements   by 

this   view   is   neither  to  initiate  rhythm  pro-      ;  .-;,  -^ 

duction   nor   control   the  orderly  sequence  of     •  ? 

the  beats  but  to  regulate  the  whole  organ  as 
regards  its  four  main  functions,  namely,  rhythm, 
conduction,  energy  production,  and  irritability. 
The  details  of  this  regulation  are  discussed 
later.  ^llW^^^^^f^  Hv 

Relation  of  Rhythm  to  Nutrition.—  The 
whole  heart,  like  the  muscular  parts  of  which 
it  is  composed,  responds  delicately  to  its  con- 
dition  of  nutrition.  In  the  frog  and  the  turtle 

hearts  the  muscular  fibers  are  brought  in  inti-  ^  „• 

mate  contact  with  the  blood  contained  within      r  ? 

the  cavities.     In  the  mammalian  heart,  on  the 

- 

other  hand,  a  distinct  system  of  vessels,  the       r^OT$^i£? 

coronary  vessels  and  the  vessels  of  Thebesius,        ;3S§^^      ^P^:      ^ 

supply  blood  to  the  walls.     If  the  heart  is  sup-        ^Jjjt'V          ^£ 

plied  with  nutrient  fluid  similar  to  its  normal        >igjf       ;. 

blood,  and  with  proper  aeration  to  insure  plenty 

of  oxygen,  it  contracts  with  a  strong  rhythm  for 

many  hours.     This  rhythm,  however,  responds 

quickly  to  changes  in  the  composition  of  the      '"  ;;"c^:fg  "..- 

nutrient  fluid.     An  abundant  supply  of  oxygen  :L  " 

is  absolutely  necessary  to  the  maintenance  of 

rhythm  in  the  mammalian  heart,  though  the        !p  £F:?"--  .^B         "c 


heart,  especially  a  cold-blooded  heart,  will  con-      vj.  ^x-.  -  :      '.?,.*•  <i- 

tract  for  a  time  in  an  atmosphere  of  hydrogen.      ^^^.^          ;^  ^      K""1 

No  doubt  the  organic  constituents  of  blood  are      '^^fe^'         %> 

very   essential   to  the  prolonged  maintenance 

of  rhythm  in  the  heart,  but  the  heart  is  not 

dependent  on  these  ingredients  for  its  stimulus 


208 


THE    CIRCULATION    OF   THE   BLOOD 


production.  The  inorganic  salts  seem  to  be  peculiarly  closely  related 
to  the  development  and  character  of  the  cardiac  rhythm,  figures  171,  172, 
and  173.  Both  the  cold-blooded  heart  and  the  mammalian  heart  respond 
very  quickly  to  the  influence  of  these  salts.  The  details  of  this  influence 
have  been  discussed  on  page  199.  It  is  somewhat  surprising,  however, 
that  the  highly  organized  mammalian  heart  will  contract  rhythmically 
for  hours  on  purely  inorganic  nutrient  fluid,  provided  only  that  the  oxygen 
be  supplied  in  sufficient  quantity  and  under  high  enough  tension.  The 
isolated  mammalian  heart  also  responds  sharply  to  a  change  in  the  salt 
content  of  the  perfusing  solution.  For  example,  addition  of  potassium 
chloride  to  a  Locke  solution  slows  or  even  suppresses  the  rate,  as  is  shown 
in  figure  178. 

Irregularities  of  Cardiac  Rhythm. — There  are  a  number  of  cardiac 
irregularities  in  rhythm  that  are  due  to  variations  in  irritability,  con- 
ductivity, or  other  of  the  normal  properties  of  the  differentiated  tissue  of 
the  heart  itself.  The  phenomena  of  this  type  of  most  common  occurrence 
are  heart  block,  extra  ventricular  systoles,  and  auricular  fibrillation. 

Heart  Block. — It  occasionally  happens  that  the  heart  rate  becomes 
very  slow,  30  or  40  a  minute,  and  the  rate  does  not  vary  much  from  this 


FIG.  178(1. — An  electrocardiogram,  lead  II,  and  auricular  and  ventricular  muscle 
tracings,  A  and  V  taken  simultaneously  from  the  dog's  heart.  Two  extra  ventricular 
contractions  are  artificially  produced  by  stimulation  of  the  right  ventricle.  They 
show  the  characteristic  right  ventricular  dominance.  The  first  is  taller  because  the  P 
of  the  natural  stimulus  and  the  R  of  the  artificial  coincide.  Time,  fifths  and  twenty- 
fifths  of  a  second.  (Lewis.) 

low  level.  On  careful  examination  it  is  found  that  the  rate  is  ventricular 
only  and  that  the  auricle  is  contracting  much  faster.  The  latter  may  con- 
tract in  multiples  of  the  ventricular  rate,  2-1,  3-1,  or  4-1  rhythm  in 
which  case  there  is  said  to  be  partial  block.  Only  every  second,  third, 
etc.,  contraction  reaches  the  ventricle.  Or  the  two  rates  may  be  wholly 
independent  as  in  total  block. 


IRREGULARITIES    OF    CARDIAC    RHYTHM 


209 


The  conducting  bundle  or  path  between  the  sino-auricular  node  where 
the  normal  beat  arises  and  the  auriculo-ventricular  node  is  usually  at 
fault  in  partial  block.  In  complete  block  the  independent  ventricular 
beats  start  from  a  rhythmic  center  in  the  auriculo-ventricular  node  as  a 
rule.  But  the. block  may  be  in  the  His  bundle  itself  in  which  case  the 
rhythm  production  is  low  in  the  conducting  system  or  even  in  the  ven- 
tricular muscle. 

Extra  Ventricular  Systoles. — When  for  any  reason  some  portion  of  the 
ventricular  complex  becomes  excessively  irritable  its  rhythmicity  may 
be  so  much  increased  that  it  starts  an  independent  contraction  before  the 
normal  ventricular  stimulus  reaches  the  muscle.  This  leads  to  a  con- 
traction with  the  shortened  period  between  beats  and  is  called  an  extra- 
ventricular  systole.  As  a  rule  there  is  a  longer  or  compensatory  pause 
following  an  extra  systole  after  which  the  regular  rhythm  again  becomes 
dominant.  Most  people  of  middle  or  old  age  experience  occasional  extra- 
systoles.  In  disease  they  may  become  frequent  and  troublesome  but 
they  are  of  no  particular  importance. 

Auricular  Flutter  and  Fibrillation. — A  type  of  irregularity  that  is  more 
common  is  that  of  an  auricular  rate  far  above  the  normal,  i.e.,  150  or  even 
more  a  minute.  These  are  due  to  hyperirritability  of  the  auricle  often 


FIG.  i  j8b. — Electrocardiogram  of  auricular  fibrillation. 

with  an  ectopic  center  of  rhythm  production.  When  such  hearts  are 
examined  with  the  electrocardiograph  or  with  the  polygraph  it  becomes 
evident  that,  the  auricle  is  contracting  at  a  very  much  higher  rate  than  the 
ventricle.  The  ventricle  does  not  respond  to  every  auricular  contraction, 
i.e.,  in  this  kind  of  block  the  stimulus  falls  within  the  refractory  period  of 
the  ventricles.  Each  auricular  contraction  is  complete  and  the  series  is 
regular  but  the  rate  is  above  that  which  can  be  conducted  to  the  ventricle. 
Auricular  flutter  is  the  term  that  describes  this  phenomenon. 

The  auricular  contractions  are  not  always  coordinated  and  com- 
plete. They  sometimes  begin  in  apparently  many  foci  at  once  so  that 
no  rhythmic  center  controls  the  entire  auricular  muscle.  Such  contrac- 
tions are  called  fibrillation.  In  auricular  fibrillation  individual  muscle 


210  THE    CIRCULATION   OF   THE   BLOOD 

cells  may  contract  and  relax  at  as  high  as  400  a  minute.  These  con- 
tractions spread  to  the  conducting  bundle  system  at  irregular  intervals 
and  the  ventricle  contracts  at  unevenly  spaced  intervals.  The  arterial 
pulse  beats  are  also  irregularly  spaced,  figure  1 786.  When  ventricular  fibril- 
lation occurs  it  quickly  produces  death,  but  auricular  fibrillation  only 
reduces  the  efficiency  of  filling  the  ventricle.  The  physician  controls 
auricular  fibrillation  by  reducing  the  irritability  of  the  auricle  and  the 
conductivity  of  the  bundle  system. 

THE  REGULATIVE  INFLUENCE  OF  THE  CENTRAL  NERVOUS 
SYSTEM  ON  THE  HEART. 

The.heart  is  capable  of  automatic  rhythmic  movement,  yet  while  in  the 
body  its  beats  are  under  the  constant  control  of  the  central  nervous  system. 
The  influence  which  is  exerted  by  the  central  nervous  system  is  of  two 
kinds:  first,  in  the  direction  of  slowing  or  inhibiting  the  beats,  and  second, 


FIG.  1 79.— Effect  on  the  Heart  Rate  and  on  the  Arterial  Blood  Pressure  of  Stimu- 
lating the  Right  Vagus  of  the  Dog.  Stimulus  applied  at  the  mark  "on"  and  removed 
at  "off."  Pressure  in  millimeters  of  mercury  shown  by  the  scale  to  the  left.  Time  in 
seconds.  (Hill  and  Chilton.) 

in  the  direction  of  accelerating  or  augmenting  the  beats.  The  influence 
of  the  first  kind  is  brought  to  bear  upon  the  heart  through  the  fibers  of  the 
pneumogastric  or  vagus  nerves,  and  that  of  the  second  kind  through  the 
sympathetic  nerves. 

The  Inhibitory  Nerves. — It  has  long  been  known,  indeed  since  the 
experiments  of  the  Weber  brothers  in  1845,  that  stimulation  of  one  or 


THE    INHIBITORY    NERVES  211 

both  vagi  produces  slowing  of  the  rhythm  of  the  heart.  It  has  since  been 
shown,  in  all  of  the  higher  vertebrate  animals  experimented  with,  that 
this  is  the  normal  reaction  to  vagus  stimulation.  Moreover,  a  section 
of  one  vagus,  or  at  any  rate  of  both  vagi,  produces  acceleration  of  the 


FIG.  1 80. — Tracing  Showing  Action  of  the  Vagus  on  the  Heart  of  the  Terrapin.  A  ur, 
Auricular;  vent,  ventricular  tracing.  The  part  between  words  "on"  and  "off"  indicates 
a  period  of  vagus  stimulation.  The  part  of  tracing  to  the  left  shows  the  regular  contrac- 
tions before  stimulation.  During  stimulation,  and  for  some  time  after,  the  beats  of  the 
auricle  and  ventricle  are  arrested.  After  they  commence  again  the  auricle  contracts 
weakly  at  first,  but  soon  acquires  a  much  greater  amplitude.  The  ventricular  contrac- 
tions that  follow  the  first  weak  auricular  contractions  are  maximal  in  the  terrapin,  but 
not  so  in  the  frog.  See  next  figure.  Time  in  seconds.  (Carr.) 

pulse  by  breaking  the  pathway  from  the  vagus  center  to  the  heart; 
stimulation  of  the  distal  or  peripheral  end  of  the  divided  nerve  normally 
produces  slowing  or  stopping  of  the  heart  beats,  showing  that  the  fibers 
are  efferent  and  thus  carry  the  nerve  impulses  toward  the  heart. 


FIG.  181— Tracing  Showing  Diminished  Amplitude  and  Slowing  of  the  Pulsations  of 
the  Auricle  and  Ventricle  of  the  frog  without  Complete  Stoppage  during  Stimulation  of 
the  Vagus.  (Gaskell.) 

It  appears  that  any  kind  of  stimulus,  either  chemical,  mechanical,  elec- 
trical, or  thermal,  produces  the  same  effect,  but  that  of  these  the  most 
potent  is  a  rapidly  interrupted  induction  current.  A  certain  amount  of 
confusion  has  arisen  as  to  the  effects  of  vagus  stimulation  in  consequence 
of  the  fact  that  fibers  of  the  sympathetic  nerve  run  within  the  trunk  of  the 


212  THE    CIRCULATION    OF   THE   BLOOD 

vagus  nerves  of  some  animals,  for  example,  the  frog.  Speaking  generally, 
however,  excitation  of  any  part  of  the  trunk  of  the  vagus  produces  inhibi- 
tion, the  stimulus  being  particularly  potent  if  applied  to  the  points  where 
the  nerves  enter  the  substance  of  the  heart  at  the  situation  of  the  sinus 
ganglia.  The  stimulus  may  be  applied  to  either  vagus  with  like  effect. 
There  are  quantitative  differences,  however,  between  the  right  and  left 
vagi.  The  right  vagus  usually  has  the  greater  effect  on  rhythm. 

The  effect  of  the  stimulation  of  the  vagus  is  threefold — to  slow  the  rate, 
or  even  to  bring  the  heart  to  a  complete  standstill,  to  produce  a  decrease 
in  the  amplitude,  and  to  delay  conduction  through  the  bundle  system. 
The  slowing  does  not  take  place  until  after  the  lapse  of  a  short  latent 
period  during  which  one  or  more  contractions  may  occur.  The  stoppage 
may  be  due  either  to  prolongation  of  the  diastole  or  to  diminution  of  the 
systole.  Vagus  stimulation  inhibits  the  spontaneous  beats  of  the  heart 
only,  it  does  not  entirely  suppress  the  irritability  of  the  heart  muscle, 
since  mechanical  stimulation  may  bring  out  a  beat  during  the  pause 
caused  by  vagus  stimulation.  The  inhibition  of  the  beats  varies  in 
duration  according  to  the  strength  of  the  stimulus  and  the  animal  stimu- 
lated. The  heart  of  the  terrapin  can  be  completely  inhibited  for  hours 
with  a  strong  stimulus.  This  phenomenon  is  shown  in  figure  180,  which 
illustrates  the  action  of  the  vagus  on  the  terrapin's  heart. 

The  heart  of  a  dog  escapes  from  complete  inhibition  in  a  few  seconds. 
When  the  beats  reappear,  the  first  few  are  usually  feeble,  after  a  time  the 
contractions  become  more  and  more  strong,  and  may  soon  exceed  both  in 
amplitude  and  frequency  those  which  occurred  before  the  application  of 
the  stimulus.  If  the  stimulation  is  prolonged,  the  inhibition  escapes 
to  a  slow  rate,  much  under  the  normal  rate.  It  is  held  there  with  some 
variations  until  the  stimulus  ceases.  This  is  due  to  the  fact  that,  in  the 
dog  at  least,  the  stimulation  reacts  more  strongly  on  rhythm  production 
at  the  sino-auricular  nodal  center,  holding  it  in  check  with  a  strength  that 
does  not  inhibit  the  auriculo-ventricular  nodal  rhythm.  The  funda- 
mental rhythm  of  the  latter  center  is  at  a  slower  rate.  The  escape  is  to  the 
auriculo-ventricular  nodal  rhythm. 

The  inhibitory  fibers  have  their  origin  in  nerve  cells  in  the  nucleus  of 
the  vagus,  and  of  the  glosso-pharyngeal,  located  in  the  floor  of  the  fourth 
ventricle.  These  cells  have  not  been  exactly  identified,  but  the  center  is 
called  the  cardio-inhibitory  center.  The  center  is  a  bilateral  one  and  the 
fibers  from  it  pass  into  the  great  vagus  trunk  to  be  distributed  to  the  heart 
through  superior  and  inferior  cardiac  branches  which  help  to  form  the 
cardiac  plexus.  Within  the  heart  the  inhibitory  fibers  form  synapses  with 
cells  whose  axones  reach  the  cardiac  muscle  cells.  The  cardiac-inhibitory 
center  is  in  more  or  less  constant  tonic  activity,  and  the  tonic  influence  is 
eliminated  when  both  nerves  are  cut,  figure  182. 


THE    ACCELERATOR    NERVES 


213 


Inhibitory  Reflexes. — The  inhibitory  center  is  influenced  by  afferent 
nerve  impulses  which  may  reach  it  from  the  heart  itself  by  the  depressor 
nerve,  or  from  other  parts  of  the  body.  These  reflex  stimulations  of 
the  vagus  center  are  constantly  occurring  during  our  daily  life  and  are 
the  most  potent  factors  in  co-ordinations  going  on  between  the  heart  and 
the  rest  of  the  body. 


FIG.  182. — Arterial  Blood  Pressure  of  the  Dog,  Showing  the  Effect  on  the  Heart  Rate 
of  Cutting  both  Vagus  Nerves  as  marked.  The  scale  to  the  left  shows  the  pressure  in 
millimeters  of  mercury.  Time  in  seconds.  The  momentary  inhibition  just  before  the 
nerves  were  cut  is  probably  due  to  mechanical  stimulation  of  the  nerves.  (Hill  and 
Chilton.) 

The  vagus  trunk  itself  contains  afferent  fibers,  the  depressor  nerves, 
that  arise  from  sensory  endothelia  in  the  heart  itself  and  in  the  aortic 
arch.  These  endings  are  stimulated  by  excessive  mechanical  pressure. - 
Their  nerve  impulses  react  on  the  vagal  motor  cells  to  produce  reflex 
inhibition,  hence  the  cardiac  slowing  that  relieves  the  pressure  that  pro- 
duced the  reflex.  This  reflex  apparatus  is  one  of  the  most  interesting 
self  protecting  mechanisms  in  the  mammalian  body. 

Rhythmical  alterations  of  the  heart  rate  occur  in  association  with  the 
effects  of  the  mechanical  variations  of  pressure  of  the  thorax  on  the  heart 
and  blood  vessels.  Apparently  the  cardio-inhibitory  center  is  stimulated 
during  the  rise  of  blood  pressure.  The  activity  of  the  center  produces  a 
slower  rate  of  the  heart  during  expiration,  shown  in  figure  243.  This 
variation  in  heart  rate  disappears  when  the  vagi  are  cut  off  from  the 
center.  The  variations  from  this  cause  are  called  sinus  arrhythmia  in 
clinical  literature.  Such  variations  are  purely  physiological  and  normal 
in  character. 

The  Accelerator  Nerves. — The  influence  of  the  accelerator  nerves 
distributed  to  the  heart  through  the  thoracic  sympathetic,  is  the  reverse 
of  that  of  the  vagus.  Stimulation  of  the  sympathetic,  even  of  one  side, 
produces  acceleration  of  the  rate  of  the  heart-beats,  augmentation  of  the 
amplitude,  or  force,  and  better  or  at  least  faster  conduction  through  the 
nodal  system  according  to  certain  observers.  Section  of  the  nerveproduces 


214 


THE    CIRCULATION    OF   THE   BLOOD 


slowing.  The  action  of  the  nerve  is  more  properly  termed  augmentor. 
The  sympathetic  or  augmentor  differs  from  the  vagus  in  several  particulars. 
First,  the  stimulus  required  to  produce  any  effect  must  be  more  powerful 
than  is  the  case  with  vagus  stimulation.  Second,  a  longer  time  elapses 
before  the  effect  is  manifest.  Third,  the  augmentation  is  followed  by 
exhaustion,  the  beats  becoming  after  a  time  feeble  and  less  frequent. 


FIG.  183. — Diagrammatic  Representation  of  the  Origin  and  Course  of  the  Cardiac 
Nerves  in  the  Dog,  showing  the  Constituent  Neurones.  D  1-5,  First  to  fifth  dorsal  spinal 
nerves.  Inhibitory  fibers  in  blue,  accelerators  in  red.  (Modified  from  Moret.) 

The  fibers  of  the  sympathetic  system,  which  influence  the  heart-beat  in 
the  frog,  leave  the  spinal  cord  by  the  anterior  root  of  the  third  spinal 
nerve.  They  pass  by  the  ramus  communicans  to  the  third  sympathetic 
ganglion,  thence  to  the  second  ganglion,  the  annulus  or  ansa  (around  the 
subclavian  artery),  through  the  first  ganglion,  and  along  the  main  trunk 
to  the  exit  of  the  vagus  from  the  cranium.  There  the  two  nerves  join 


THE    ACCELERATOR    NERVES 

and  run  down  to  the  heart  within  a  common  sheath,  forming  the  vago- 
sympathetic  trunk.  Stimulation  of  the  accelerators  of  the  frog  must  be 
applied  to  the  pathway  before  the  fibers  join  the  common  trunk  if  uncom- 
plicated augmentation  is  to  be  secured.  On  stimulation  of  the  mixed 
vago-sympathetic  trunk  inhibition  ordinarily  occurs  at  once.  Augmentor 
effects  come  on  only  after  the  inhibition  has  disappeared,  usually  fifteen 
or  twenty  seconds  later.  If  the  vagal  influence  is  first  removed  by  a 
specific  poison,  atropine,  then  on  stimulation  pure  augmentation  results 
at  once.  This  method  applied  to  the  frog  is  one  of  the  most  satisfactory 
methods  of  illustrating  the  different  elements  in  cardiac  augmentation. 

In  the  dog  the  augmentor  fibers  leave  the  cord  by  the  anterior  roots  of 
the  second  and  third  dorsal  nerves,  and  possibly  also  by  the  first,  fourth, 
and  fifth  dorsal  nerves.  They  pass  by  the  rami  communicantes  to  the  gang- 
lion stellatum,  or  first  thoracic  ganglion,  and  around  the  ansa  to  the  inferior 
cervical  ganglion  of  the  sympathetic.  Fibers  from  the  ansa  or  from  the 
inferior  cervical  ganglion  proceed  to  the  heart,  figure  183.  The  course  of 
the  augmentor  fibers  in  the  spinal  cord  is  not  so  well  known  except  that 
they  originate  in  an  augmentor  center  in  the  medulla.  The  circulation 
of  venous  blood  appears  to  stimulate  the  augmentor  center,  and  of  highly 
oxygenated  blood  the  inhibitory  center. 

The  accelerator  center,  like  the  inhibitory,  is  in  constant  tonic  activity; 
and  the  cardiac  acceleration  on  cutting  the  vagi,  shown  in  figure  182,  is  in 
part  to  be  ascribed  to  this  tone.  When  both  nerves  are  stimulated 
together,  the  resulting  rate  is  the  algebraic  sum  of  the  opposed  influences, 
according  to  Hunt.  The  accelerator  center  is  influenced  by  afferent 
impulses  arising  throughout  the  body,  and  these  reflexes  contribute  to  the 
general  co-ordination  of  the  chest  with  the  activities  of  the  body. 

In  addition  to  direct  and  reflex  stimulation,  impulses  passing  down  from 
the  cerebrum  may  have  a  similar  effect,  psychic  stimulation. 

Other  Influences  which  Affect  the  Heart. — A  great  variety  of  special 
conditions  influence  the  heart's  action  in  the  normal  body,  conditions  that 
are  not  discussed  directly  under  any  of  the  categories  treated  above.  Of 
these  may  be  mentioned  the  coronary  circulation,  temperature,  mechanical 
tension,  age. 

The  Coronary  Circulation. — The  contractions  of  the  heart  cannot  long 
be  maintained  without  a  due  supply  of  blood  or  other  nutrient  fluid.  The 
nutrient  fluid  for  the  heart  of  man  and  the  mammals  is  supplied  from  the 
coronary  arteries  and  the  vessels  of  Thebesius.  The  coronary  arteries  arise 
from  the  base  of  the  aorta,  where  they  receive  the  benefit  of  the  highest 
arterial  pressure.  The  coronary  arteries  are  terminal  arteries;  that  is, 
they  do  not  permit  the  establishment  of  a  collateral  circulation  when  one 
of  their  branches  is  blocked.  If  the  block  be  complete,  that  portion  of  the 
heart  wall  supplied  by  the  branch  dies.  The  immediate  effect  of  the 


2l6  THE    CIRCULATION    OF   THE   BLOOD 

closure  of  a  large  coronary  branch,  in  the  dog,  may  be  occasional  and 
transient  irregularity  or  arrest  of  the  ventricular  contractions  preceded 
by  irregularities  in  the  force  of  the  contractions  and  a  diminution  in  the 
amount  of  work  performed.  The  force,  rather  than  the  rate,  of  the  ven- 
tricular contractions  is  closely  dependent  upon  the  blood  supply  to  the 
coronary  arteries.  Porter  and  others  have  shown  that  the  pressure  in  the 
coronary  vessels  follows  closely  the  pressure  in  the  aorta  and  that  there  is 
not,  as  formerly  claimed,  a  closure  of  these  vessels  by  the  pressure  of  the 
systole  of  the  ventricle. 

The  vessels  of  Thebesius,  which  have  been  demonstrated  to  open  both 
into  the  auricular  and  ventricular  cavities,  must  now  be  looked  upon,  ac- 
cording to  the  investigations  of  Pratt,  as  an  important  source  of  cardiac 
nutrition.  Blood  may  pass  through  them  by  way  of  connecting  branches 
to  the  coronary  arteries  and  veins.  Pratt  succeeded  in  maintaining  cardiac 
contractions  for  several  hours  when  the  only  source  of  nutrition  was  from 
these  vessels.  This  source  of  nutrition  may  account  for  the  survival  of 
hearts  for  years  where  pronounced  arterio- sclerosis  of  the  coronary  arteries 
exists. 

Alteration  of  Temperature. — The  effect  of  cold  is  to  slow  the  rate  of  the 
heart-beat.  If  the  heart  of  a  frog  be  cooled  down  to  o°  C.  it  will  stop 
beating,  but  when  the  temperature  of  the  surrounding  lymph  or  blood  is 
again  raised,  it  will  renew  its  spontaneous  beats.  The  effect  of  heat  is  to 
quicken  and  shorten  the  heart-beats,  but  at  a  moderate  temperature,  20° 
C.,  the  contractions  are  increased  in  force,  figure  177. 

The  isolated  mammalian  heart  is  influenced  by  temperature  variations 
in  much  the  same  way  as  that  of  the  frog.  It  will  contract  slowly  in  a  low 
temperature  and  rapidly  in  a  temperature  higher  than  that  normal  to  the 
body.  The  very  rapid  heart  in  some  high  fevers  is  in  part  due  to  the  increase 
in  temperatures  which  affects  the  heart  directly. 

Mechanical  Tension. — The  mechanical  factors  produced  by  the  heart- 
beat are  so  prominent  that  it  would  be  surprising  indeed  if  there  were  no 
reaction  of  these  mechanical  conditions  on  the  heart  itself.  The  isolated 
cardiac  muscle  responds  very  quickly  to  variations  in  tension.  Beginning 
with  a  low  tension  the  activity  of  heart  muscle  is  increased  up  to  a  certain 
optimum  tension,  after  which  further  increase  is  unfavorable  to  the  develop- 
ment of  automatic  rhythm.  A  quite  strong  stretching  will  paralyze  the 
muscle. 

Tension  on  the  whole  heart  influences  its  activity,  not  only  through  the 
effects  on  the  muscle,  but  indirectly  through  the  nervous  mechanism.  High 
tension,  such  as  contracting  against  a  high  aortic  pressure,  stimulates  sensory 
nerves  of  the  heart  which,  acting  through  the  depressor  nerve  on  the  inhibi- 
tory center,  produce  reflex  slowing  of  the  heart.  It  also  produces  reflex 
vaso-dilatation.  Both  reflexes  relieve  the  high  tension  on  the  heart.  This 


ALTERATION  OF  TEMPERATURE  217 

nerve  reaction  takes  place  with  a  tension  which  still  mechanically  stimulates 
the  cardiac-muscle  substance,  and  the  inhibitory  effects  must  therefore  over- 
come the  direct  stimulating  effect  of  the  tension  on  the  muscle  fibers. 

Age,  Sex,  etc. — The  average  heart  rate  for  the  normal  adult  man  is  72 
times  a  minute,  but  this  rate  will  vary  much  in  different  individuals  accord- 
ing to  the  age,  sex,  size,  and  personal  equation.  The  frequency  of  the  heart's 
action  gradually  diminishes  from  the  commencement  to  near  the  end  of  life, 
but  is  said  to  increase  again  somewhat  in  extreme  old  age,  thus: 

Before  birth  the  average  number  of  pulsations 

per  minute  is 1 50 

Just  after  birth 130  to  140 

During  the  first  year 1 1 5  to  130 

During  the  second  year 100  to  1 1 5 

During  the  third  year 90  to  100 

About  the  seventh  year 85  to     90 

About  the  fourteenth  year 80  to     85 

In  adult  age.  .    70  to     80 

In  old  age 60  to     70 

In  decrepitude 65  to     75 

The  heart  rate  is  greater  in  woman  than  in  man.  It  is  also  greater  in 
small  than  in  large  individuals.  The  rate  varies  from  the  type  in  certain 
individuals  where  no  cause  can  be  assigned  other  than  personal  equation. 

Poisons  and  Other  Chemical  Substances. — A  large  number  of  chemical 
substances  have  a  distinct  effect  upon  the  cardiac  contractions.  Of  these 
the  most  important  are  atropine,  muscarine,  digitalis,  barium,  nicotine, 
caffeine,  etc. 

Atropine  produces  considerable  augmentation  of  the  heart-rate,  and 
when  acting  upon  the  heart  prevents  inhibition  by  vagus  stimulation.  Its 
effects  are  produced  by  poisoning  the  nerve  endings  of  the  vagus  within 
the  heart.  When  these  endings  are  poisoned  stimuli  arising  in  the  inhibi- 
tory center  of  the  medulla  (tonic  activity),  or  artificially  applied  to  the 
vagus,  cannot  reach  the  heart  muscle,  and  inhibition  is  impossible. 

Muscarine,  which  is  obtained  from  various  species  of  poisonous  fungi, 
produces  marked  slowing  of  the  heart-beats,  and,  in  larger  doses,  stoppage 
of  the  heart.  It  produces  an  effect  similar  to  that  of  prolonged  vagus  stimu- 
lation. The  effect  can  be  removed  by  the  action  of  atropine,  hence  is 
supposed  to  stimulate  the  nerve  endings  of  the  vagus. 

Digitalis  slows  the  heart  by  stimulating  the  vagi  at  their  origin  in  the 
inhibitory  center  in  the  medulla.  The  heart  muscle  itself  is  also  rendered 
more  excitable. 

Veratrine  and  aconitine  have  a  somewhat  similar  effect. 

Nicotine  and  caffeine  are  both  very  powerful  cardiac  stimulants.  The 
great  injurious  effects  of  nicotine  on  the  heart  are  due  to  two  causes,  first, 


21 8  THE    CIRCULATION   OF   THE   BLOOD 

to  paralysis  of  the  nervous  mechanism  and  relative  loss  of  control, 
second,  to  the  great  direct  stimulation  of  the  cardiac  muscle.  The 
constant  overuse  of  tobacco,  therefore,  very  sharply  weakens  the  effi- 
ciency of  the  heart.  Caffeine  does  not  lead  to  so  great  disturbance  of 

the  heart's  nutrition  as  does  nicotine. 

t 

THE  CIRCULATION  THROUGH  THE  BLOOD  VESSELS. 

Blood-Pressure. — The  subject  of  blood-pressure  has  been  already 
incidentally  mentioned  more  than  once  in  the  preceding  pages;  the  time  has 
now  arrived  for  it  to  receive  more  detailed  consideration. 

That  the  blood  exercises  pressure  upon  the  walls  of  the  vessels  containing 
it  is  due  to  the  following  facts: 

The  heart  at  each  contraction  forcibly  injects  a  considerable  amount  of 
blood,  80  to  100  c.c.,  suddenly  and  quickly  into  the  arteries. 

The  arteries  are  highly  distensible  and  stretch  to  accommodate  the  extra 
amount  of  blood  forced  into  them.  The  arteries  are  already  full  of  blood 
at  the  commencement  of  the  ventricular  systole,  since  there  is  not  sufficient 
time  between  the  heart-beats  for  the  blood  to  pass  into  the  veins. 

There  is  a  distinct  resistance  interposed  to  the  passage  of  the  blood  from 
the  arteries  into  the  veins  by  the  enormous  number  of  minute  vessels,  small 
arteries  (arterioles)  and  capillaries,  into  which  the  main  artery  has  been 
ultimately  broken  up.  The  sectional  area  of  the  capillaries  is  several  hun- 
dred times  that  of  the  aorta,  and  the  friction  generated  by  the  passage  of 
the  blood  through  these  minute  channels  opposes  a  considerable  hindrance 
or  resistance  in  its  course.  The  resistance  thus  set  up  is  called  peripheral 
resistance.  The  friction  is  greater  in  the  arterioles,  where  the  current  is 
comparatively  rapid,  than  in  the  capillaries,  where  it  is  slow. 

The  interaction  of  these  factors — heart-beat,  elastic  vessels,  and  periph- 
eral resistance — is  sufficient  to  maintain  a  flow  of  blood  through  the  entire 
circulatory  system.  It  is  the  interrelation  of  these  factors  which  main- 
tains an  even  and  steady  flow  through  the  capillaries  and  past  the  tissues, 
where  it  is  desirable  that  the  conditions  of  blood  flow  should  be  most  con- 
stant if  the  purposes  of  nutrition  are  to  be  best  accomplished.  In  fact, 
we  shall  even  find  that  it  is  the  interaction  of  these  same  factors,  together 
with  the  possibility  of  variations  through  regulation  by  their  nerve-motor 
mechanisms,  that  we  have  the  great  variations  and  adjustments  of  blood- 
pressure,  speed  of  flow,  volume  of  flow,  and  the  regulation  of  volume  in 
particular  parts  of  the  body  or  local  control. 

Arterial  Blood -Pressure. — That  the  blood  exerts  considerable  pres- 
sure upon  the  arterial  walls  in  keeping  them  in  a  stretched  or  distended 
condition  may  be  readily  shown  by  puncturing  any  artery.  The  blood  is 
instantly  projected  with  great  force  through  the  opening,  and  the  jet  rises 
to  a  considerable  height,  the  exact  level  of  which  varies  with  the  size  of  the 


THE    CIRCULATION    THROUGH    THE    BLOOD   VESSELS  2IQ 

artery  experimented  upon.  If  a  large  artery  be  punctured  the  blood  may 
be  projected  upward  for  several  feet,  whereas  if  it  is  a  small  artery  the  jet 
does  not  rise  so  high.  Another  characteristic  of  the  jet  of  blood  from  a  cut 
artery,  particularly  well  marked  if  the  vessel  be  a  large  one  and  near  the 
heart,  is  the  intermittent  character  of  the  outflow.  If  the  artery  be  cut 
across,  the  jet  issues  with  force,  chiefly  from  the  central  end.  If  there  is 
considerable  anastomosis  of  vessels  in  the  neighborhood  the  jet  from  the 
peripheral  end  may  be  almost  as  forcible  and  as  intermittent  as  that  from 
the  central  end.  The  intermittent  flow  in  the  arteries  due  to  the  action 
of  the  heart,  and  which  represents  the  systolic  and  diastolic  alterations  of 
blood-pressure,  may  be  felt  if  the  finger  be  placed  upon  a  sufficiently 
superficial  artery.  The  finger  is  apparently  raised  and  lowered  by  the 
intermittent  distention  of  the  vessel  occurring  at  each  heart-beat.  This 
intermittent  distention  of  the  artery  is  what  is  known  as  the  pulse,  to  the 
further  consideration  of  which  we  shall  presently  return,  but  we  may  say 
here  that  in  the  normal  condition  the  pulse  is  a  characteristic  of  the  arterial, 
and  is  absent  from  the  venous,  flow. 

At  the  same  time  it  must  be  recollected  that  in  the  veins  also  the  blood 
exercises  a  pressure  on  the  containing  vessels,  though  it  is  small  when 
compared  with  the  arterial  pressure.  As  might  be  expected,  therefore, 
the  blood  is  not  expelled  with  so  much  force  if  a  vein  be  punctured  or  cut. 
The  flow  from  the  cut  vein  is  continuous  and  not  intermittent,  and  the 
greater  amount  of  blood  comes  from  the  peripheral  and  not  from  the 
central  end,  as  is  the  case  when  an  artery  is  severed. 

Methods  of  Measuring  Arterial  Blood -Pressure. — The  pressure  in 
an  artery  may  be  measured  by  cutting  the  vessel  and  introducing  into  it  a 
cannula  and  connecting  the  cannula  with  a  tall  vertical  glass  tube.  When  the 
blood  in  the  vessel  is  released  to  the  cannula,  a  column  of  blood  will  rise  in  the 
tube  at  once  to  the  height  that  can  be  supported  by  thfe  pressure  in  that  par- 
ticular vessel.  If  the  vessel  be  an  artery,  the  blood  will  rise  several  feet, 
according  to  the  distance  of  the  vessel  from  the  heart,  and  when  the  pressure 
has  reached  its  highest  point  it  will  be  seen  to  oscillate  with  the  heart-beats. 
This  experiment  shows  that  the  pressure  which  the  blood  exerts  upon  the 
walls  of  the  containing  artery  equals  the  pressure  of  a  column  of  blood  of  a 
certain  height.  In  the  case  of  the  rabbit's  carotid  it  is  equal  to  90  to  120  cm. 
of  blood,  or  rather  more  than  the  same  height  of  water.  In  the  case  of  the 
vein,  if  a  similar  experiment  be  performed,  blood  will  rise  in  the  tube  only 
for  8  or  10  cm.  or  less. 

The  usual  method  of  estimating  the  amount  of  blood-pressure  differs 
somewhat  from  the  foregoing  simple  experiment.  Instead  of  a  simple 
straight  tube  or  glass  manometer  for  measuring  the  pressure,  a  U-shaped  tube 
containing  mercury,  the  mercury  manometer,  is  employed.  The  artery  is 
connected  with  the  manometer  by  means  of  the  cannula  inserted  into  the 
vessel  as  before,  an  arrangement  being  made  whereby  the  cannula,  tubes,  etc., 


22O  THE    CIRCULATION    OF    THE   BLOOD 

are  first  filled  with  a  half -saturated  solution  of  magnesium  sulphate  or  other 
saline  to  prevent  the  clotting  of  blood  when  it  is  allowed  to  pass  from  the 
artery  into  the  apparatus.  The  loss  of  blood  is  prevented  during  the 
preparation  of  the  details  of  the  experiment  by  a  clamp  or  bull-dog  for- 
ceps. The  free  end  of  the  U-tube  of  mercury  contains  a  very  fine  glass 
or  metal  rod  with  a  bulb  which  floats  upon  the  surface  of  the  mercury 
and  oscillates  with  the  oscillations  of  the  mercury.  As  soon  as  there  is 
free  communication  between  the  artery  and  the  tube  of  mercury,  the  blood 
rushes  out  and  pushes  before  it  the  column  of  mercury.  The  mercury  will 
therefore  rise  in  the  free  limb  of  the  tube,  and  will  continue  to  do  so  until  a 
point  is  reached  which  corresponds  to  the  mean  pressure  of  the  blood-vessel 
used.  The  blood-pressure  is  thus  communicated  to  the  near  limb  of  the 
column  of  mercury;  and  the  depth  to  which  the  latter  sinks,  added  to  the 
height  to  which  it  rises  in  the  other  limb,  the  weight  of  the  saline  solution 
being  subtracted,  will  give  the  height  of  the  column  of  mercury  which  the 
blood-pressure  balances.  For  the  estimation  of  the  amount  of  blood-pressure 
one  can  make  direct  readings  at  any  given  moment  and  no  further  apparatus 


FIG.  184.  FIG.  185. 

FIG.  184. — Arterial  Cannula.  T-form  for  convenience  in  washing  out  clots. 
FIG.  185. — Ludwig's  Mercury  Manometer.  The  mercury  which  partially  fills  the 
tube  supports  a  float  in  the  form  of  a  piston,  nearly  filling  the  tube;  a  wire  is  fixed  to  the 
float,  and  the  writing  style  or  pen  is  guided  by  passing  through  the  brass  cap  of  the 
manometer  tube;  the  pressure  is  communicated  to  the  mercury  by  means  of  a  flexible 
metal  tube  filled  with  fluid. 

than  this  is  necessary.  But  in  the  more  accurate  study  of  the  variations 
of  pressure  in  the  arterial  system,  as  well  as  its  absolute  amount,  the  instru- 
ment is  usually  combined  with  a  recording  apparatus,  called  a  kymograph. 
Numerous  forms  of  recording  kymographs  are  to  be  had  in  the  market. 
These  instruments,  while  all  constructed  on  the  same  principle,  vary  chiefly 
in  the  accuracy  of  their  construction  and  convenience  of  their  adjustments. 


METHODS    OF    MEASURING    ARTERIAL   BLOOD-PRESSURE          221 

The  essential  part  of  a  recording  kymograph  consists  of  a  uniformly 
revolving  cylinder  accurately  centered  and  carrying  a  paper  on  which  a  record 
is  made  of  the  physiological  change  which  is  being  studied.  This  cylinder 
or  drum  may  be  driven  by  a  weight,  clock  spring,  electric  motor,  or  other 
mechanical  device  that  insures  uniformity  of  speed  and  which  is  capable  of 
speed  regulation.  The  cylinder  is  covered  with  glazed  paper,  blackened  in 
the  flame  of  a  lamp,  and  the  mercury  manometer  is  so  supported  that  its 
float,  provided  with  a  style,  writes  on  the  cylinder  as  it  revolves.  In  some 
of  the  instruments,  especially  Ludwig's  continuous  paper  kymograph,  a 
long  paper  band  is  made  to  pass  over  the  recording  surface  and  the  record 
itself  is  written  by  various  devices  carrying  ink. 

There  are  also  many  ways  in  which  the  mercury  manometer  may  be 
varied;  in  figure  185  is  seen  a  form  which  is  known  as  Ludwig's.  In  order 
to  obviate  the  necessity  of  a  large  quantity  of  blood  entering  the  tube  of  the 
apparatus  and  being  lost  to  the  animal,  it  is  usual  to  have  some  arrangement 
by  means  of  which  the  mercury  may,  previous  to  the  experiment,  be  forced 


wo 
to 


FIG.  186. — Tracing  of  Normal  Arterial  Pressure  in  the  Dog,  Obtained  with  the 
Mercurial  Manometer.  The  smaller  undulations  correspond  with  the  heart-beats;  the 
larger  curves  with  the  respiratory  movements.  Pressure  is  in  millimeters  of  mercury  as 
shown  by  the  scale  to  the  left.  Time  in  seconds. 

up  in  the  tube  of  the  manometer  to  the  pressure  level  corresponding  to 
approximately  the  mean  pressure  of  the  artery  experimented  with,  so  that  the 
writing  style  simply  records  the  variations  of  the  blood-pressure  above  and 
below  the  mean  pressure.  This  is  done  by  causing  the  anti-coagulant 
solution,  generally  a  saturated  solution  of  sodium  carbonate  or  of  10  per 


222  THE    CIRCULATION    OF    THE   BLOOD 

cent,  magnesium  sulphate,  to  fill  the  apparatus  from  a  bottle  suspended  at 
a  height  about  that  of  the  pressure  to  be  measured,  and  capable  of  being 
raised  or  lowered  as  required  for  the  purpose. 

The  cannula  inserted  and  tied  into  the  artery  may  be  of  several  different 
kinds.  A  glass  T-tube  with  the  end  drawn  out  and  cut  so  that  it  is  oblique, 
and  provided  with  a  slightly  constricted  neck  to  prevent  its  coming  out  of 
the  artery  easily,  is  a  very  convenient  form,  figure  184.  Of  the  two  free 
ends  of  the  T-cannula  one  is  connected  with  the  manometer,  the  other  with 
the  pressure  bottle.  The  peripheral  end  of  the  cut  artery  is  tied  to  obviate 
the  escape  of  blood.  By  this  means,  the  presssure  communicated  to  the 
column  of  mercury  is  the  forward,  and  not  the  lateral,  pressure  of  blood, 
but  there  is  very  little  if  any  difference. 

As  soon  as  the  experiment  is  begun,  the  writing  float  is  seen  to  oscillate 
in  a  regular  manner,  and  a  curve  of  blood  pressure  is  traced  upon  the  smoked 
paper  by  the  style  (or,  if  a  continuous  roll  of  unsmoked  paper  be  used,  the 
trace  is  made  by  an  inked  pen)  when  a  figure  similar  to  figure  186  will  be 
obtained.  This  indicates  two  main  variations  of  the  blood  pressure.  The 
smaller  excursions  of  the  lever  correspond  with  the  systole  and  diastole  of  the 
heart,  and  the  larger  curves  correspond  with  the  respirations,  being  called 
the  respiratory  undulations  of  blood-pressure,  to  which  attention  will  be  directed 


FIG.  187.— Tracing  of  Normal  Arterial  Pressure  Taken  from  the  Rabbit  with  a  Hurthle 
Manometer.     The  horizontal  lines  show  zero  pressure.     Time  in  seconds.     (Dreyer.) 

in  the  next  chapter.  Of  course,  the  undulations  spoken  of  are  seen  only  in 
records  of  arterial  blood-pressure.  They  are  more  clearly  marked  in  the  ar- 
teries nearer  the  heart  than  in  those  more  remote.  The  amount  of  the 
pressure  in  the  smaller  arteries  as  well  as  the  indication  of  the  systolic  rise 
of  pressure  is,  comparatively  speaking,  small. 

In  order  to  record  the  details  of  the  undulations  of  arterial  pressure,  it  is 
better  for  some  purposes  to  use  the  Hurthle  membrane  manometer  than  the 
mercurial  manometer.  Two  views  of  this  instrument  are  shown  in  figure  166. 


BLOOD-PRESSURE    MEASUREMENTS    IN    MAN  223 

The  instrument  consists  of  a  hollow  tube  and  cup  covered  with  rubber  sheet 
against  which  a  disc  supported  by  a  metal  spring  is  adjusted.  The  appara- 
tus is  filled  with  fluid,  the  interior  of  which  is  connected  with  the  artery  by 
means  of  a  metal  tube  and  cannula.  The  pressure  transmitted  to  the  appa- 
ratus tends  to  stretch  the  rubber  and  bend  the  spring,  and  the  movement  thus 
produced  is  communicated  by  means  of  a  lever  to  a  writing  style  and  so  to 
a  recording  apparatus.  This  instrument  obviates  the  errors  which  might 
be  caused  by  the  inertia  of  the  mercury  in  the  mercurial  manometer;  it  alsa 
shows  in  more  detail  the  variations  of  the  blood  pressure  in  the  vessel  during, 
and  after  each  individual  beat  of  the  heart. 

As  regards  the  actual  amount  of  blood-pressure,  from  observations  which 
have  been  made  by  means  of  the  mercurial  manometer,  it  has  been  found 
that  the  pressure  of  blood  in  the  carotid  of  a  rabbit  is  capable  of  supporting 
a  column  of  90  to  1 20  mm.  of  mercury ;  in  the  dog  i  oo  to  1 7  5  mm. ;  in  the  horse 
152  to  200  mm.;  and  in  man  the  pressure  is  estimated  to  be  about  the  same 
as  in  the  dog.  To  measure  the  absolute  amount  of  this  pressure  in  any 
artery  multiply  the  area  of  its  transverse  section  by  the  height  of  the  column 
of  mercury  which  is  already  known  to  be  supported  by  the  blood  pressure 
in  any  part  of  the  arterial  system.  The  weight  of  a  column  of  mercury  thus 
found  will  represent  the  absolute  pressure  of  the  blood.  Calculated  in  this 
way,  the  blood  pressure  in  the  human  aorta  is  equal  to  i .  93  kilogrammeters; 
that  in  the  aorta  of  the  horse  being  5  .  2  kilogrammeters;  and  that  in  the  radial 
artery  at  the  human  wrist  only  o .  08  kilogrammeter.  Supposing  the  muscu- 
lar power  of  the  right  ventricle  to  be  one-fourth  that  of  the  left,  absolute 
pressure  in  the  pulmonary  artery  will  be  only  0.5  kilogrammeter.  The 
amounts  above  stated  represent  the  arterial  tension  at  the  time  of  the 
ventricular  contraction. 

The  arterial  pressure  is  greatest  at  the  beginning  of  the  aorta,  and  de- 
creases toward  the  capillaries.  It  is  greatest  in  the  arteries  at  the  period  of 
the  ventricular  systole  and  least  during  the  diastole.  The  blood-pressure 
gradually  lessens  as  we  proceed  from  the  arteries  near  the  heart  to  those 
more  remote,  and  again  from  these  to  the  capillaries,  as  it  does  also  from 
the  capillaries  along  the  veins  to  the  right  auricle. 

Arterial  Blood -Pressure  Measurements  in  Man. — A  number  of 
instruments  have  been  devised  for  estimating  blood-pressure  in  man  for 
clinical  purposes.  Some  of  these,  though  excellent  in  principle,  are  too 
complicated  for  general  use.  The  first  simple  and  approximately  accurate 
form  of  apparatus  was  that  devised  by  Riva-Rocci  in  1896.  This  has  been 
modified  and  improved  in  minor  points  since,  but  the  principles  of  the 
original  instrument  remain  practically  the  same. 

In  brief,  the  apparatus,  figure  188,  consists  of  an  elastic  tube  ending  in  a 
rubber  bag  which  can  be  adjusted  about  the  arm,  and  a  mercury  man- 
ometer connected  with  this  tube  and  also  with  some  form  of  air  pump 


224 


THE    CIRCULATION    OF    THE    BLOOD 


used  for  inflating  the  tube  about  the  arm  and  thus  exerting  pressure  upon 
its  blood-vessels.  The  elastic  tube  is  covered  by  some  inelastic  tissue,  usu- 
ally a  leather  cuff,  in  order  that  the  inflation  of  the  bag  may  cause  the  full 
increase  of  pressure  to  be  exerted  upon  the  encased  arm.  By  inflating  the 
bag  until  the  pulse  at  the  wrist  just  disappears,  and  reading  the  height  of 
the  column  of  mercury  in  the  manometer,  the  maximum  or  systolic 
pressure  is  obtained  in  millimeters  of  mercury.  If  now  the  pressure  on 
the  arm  is  reduced  until  the  widest  oscillations  of  the  mercury  column 
are  obtained,  the  lowest  position  of  the  mercury  meniscus  represents  the 
diastolic  pressure. 

The  apparatus  depends  on  the  principle  that  an  external  pressure  just 
equal  to  the  maximal  pressure  within  an  artery  will  hold  the  vessel  in  the 
collapsed  condition,  a  fact  that  has  been  proven  for  vessels  that  are  exposed. 
An  external  pressure  that  will  just  equal  the  minimal  or  diastolic  pressure 
will  cause  a  complete  collapse  of  a  vessel  during  diastole  and  will  allow  a 
complete  expansion  of  an  artery  to  its  maximal  limits  during  the  systolic 
period  of  pressure.  In  other  words,  the  mercury  of  the  manometer  will 
oscillate  to  its  maximal.  If  the  pressure  is  reduced  to  a  still  lower  point,  it 


FIG.  188. — Riva-Rocci  Apparatus  (schematic)  for  Determining  Blood  Pressure  in  Man. 

will  not  be  sufficient  to  compress  the  artery  completely,  and  the  mercury 
oscillations  will  again  become  smaller.  In  applying  the  instrument  to  the 
brachial  artery,  one  must,  of  course,  deal  with  a  vessel  deeply  buried  in  mus- 
cular and  other  tissues.  These  latter  tissues  probably  consume  a  certain 


BLOOD-PRESSURE    MEASUREMENTS    IN   MAN 


225 


small  percentage  of  the  pressure,  an  error  which  may  be  ignored  for  all 
comparative  purposes. 

Erlanger  has  perfected  a  form  of  sphygmomanometer  which  contains  a 
very  ingenious  and  compactly  arranged  recording  device,  figure  189.  This 
instrument  has  a  mercury  manometer  from  which  the  pressures  are  read  off 
directly.  On  a  side  limb  of  the  manometer  there  is  a  rubber  bag  enclosed 
in  a  glass  bell.  The  cavity  of  the  bell  outside  of  the  rubber  bag  is  connected 


FIG.  189. — Erlanger's  Sphygmomanometer,  Shown  with  the  Rubber  Bag  Attached  to 
the  Arm.  The  picture  is  taken  at  the  end  of  an  experiment  after  the  pressure  in  the  instru- 
ment is  run  up  again  to  above  the  systolic  pressure.  The  upper  part  of  the  cylinder  shows 
a  sphygmogram  taken  with  the  instrument.  (Experiment  and  photo  by  Hill  and  Watkins.) 

with  a  recording  tambour,  the  entire  apparatus  being  fully  supplied 
with  the  necessary  valves  and  adjusting  devices  which  make  it 
mechanically  very  perfect.  The  instrument  is  mounted  on  a  stand  with  a 
small  clock  and  recording  cylinder  adapting  it  to  convenient  clinical  use. 
The  brachial  arterial  pressure  of  man  when  taken  by  this  form  of 
apparatus  has  been  found  to  vary  greatly,  but  Erlanger  gives  no  mm.  of 
mercury  as  the  average  of  observations  on  young  adults  in  the  determi- 


226  THE    CIRCULATION    OF    THE   BLOOD 

nation  of  the  systolic  pressure;  i.e.,  the  maximal  arterial  pressure.  He 
gives  for  the  diastolic  pressure  40  to  45  mm.  of  mercury  below  the  systolic 
pressure.  Other  observers  using  the  same  method  find  a  somewhat  higher 
average  pressure,  see  figure  190,  which  represents  a  fair  type  of  observation. 
The  form  of  sphygmomanometer  in  almost  universal  clinical  and 
laboratory  use  for  determining  the  arterial  blood-pressure  of  man  is  the 
aneroid  type  of  Dr.  Rogers.  This  instrument  or  its  various  modifications 
measures  the  pressure  by  means  of  the  expansion  of  an  aneroid  coupled 
with  a  mechanical  lever  and  gage  device.  The  most  widely  distributed 
forms  of  instruments  of  this  type  are  known  as  the  Tyco  and  Faught. 
These  instruments  use  an  arm  belt  and  bag  of  the  Riva-Rocci  type.  The 
rubber  bag  is  inclosed  in  a  cloth  belt  which  is  conveniently  wrapped 
around  the  arm  above  the  elbow.  The  bag  contains  two  connections 


FIG.  190. — Tracing  taken  with  Erlanger's  Sphygmomanometer.  The  figures  indicate 
pressure  in  millimeters  of  mercury.  Systolic  pressure  160;  diastolic  pressure,  120.  (New 
figure  by  Hill.) 

one  of  which  is  attached  to  the  pressure  gage,  the  other  connected  with 
a  convenient  pump  made  of  either  metal  of  rubber.  The  pressure  meas- 
urement can  be  made  directly  from  the  oscillations  of  the  dial  as  described 
for  the  Erlanger  or  Riva-Rocci  apparatus.  Readings  may  also  be  ob- 
tained by  the  palpation  of  the  artery  at  the  wrist  as  the  pulse  breaks 
through  during  gradual  reduction  of  the  pressure  in  the  arm  bag.  How- 
ever, the  most  accurate  determinations  are  made  by  the  auscultatory 
method  (Goodman  and  Howell).  A  Bowles  sphygmometroscope,  which  is 
a  stethoscope  modified  by  a  button  attached  to  the  center  of  the  disc,  is 
attached  to  the  arm  just  below  the  arm  band  at  the  inner  angle  of  the 
elbow  with  the  button  of  the  diaphragm  directly  over  the  brachial  artery 
near  its  division  into  the  ulnar  and  radial. 

In  operation  the  arm  band  is  pumped  to  a  pressure  above  that  of 
the  underlying  artery  and  then  the  pressure  very  gradually  released. 
When  the  external  pressure  just  equals  to  or  is  slightly  less  than  the 
maximum  pressure  in  the  artery,  some  fluid  will  escape  into  the  occluded 


THE    VENOUS    BLOOD-PRESSURE    AND    CAPILLARY    PRESSURE       227 


limb  of  the  artery  below  the  band.  The  flow  of  this  fluid  produces  a  very 
definite  first  sound  which  is  used  to  determine  the  moment  of  systolic 
pressure.  The  sounds  in  the  brachial  artery,  known  as  Karatkoff  sounds, 
have  been  described  as  going  through  five  phases  before  the  circulation  is 
fully  established  in  the  cut  off  artery.  The  first  phase  is  the  initial 
development  of  a  clear  cut  and  sharp  sound.  It  is  the  index  of  the 
systolic  pressure.  The  first  sound  is  followed  by  a  series  of  murmurs 
called  the  second  sound,  and  that  by  a  more  definite  tone,  the  third  phase. 
The  third  phase  will  vary  in  character  with  certain  abnormalities  in  the 
vessel  wall — thickening,  sclerosis,  etc. 

The  fourth  phase  is  the  appearance  of  a  duller  tone  of  diminishing 
intensity  which  rapidly  fades  into  no  sound,  the  so-called  fifth  phase. 
The  fourth  phase  is  taken  as  the  index  of  minimal  or  diastolic  arterial 
pressure. 

If  the  ascultatory  reading  and  the  palpation  reading  are  made  at  the 
same  time  the  latter  usually  gives  a  slightly  lower  systolic  pressure  than 
the  former.  In  other  words,  the  stethoscopic  reading  by  the  sound  is  the 
more  accurate.  The  diastolic  reading  is  far  more  accurately  determined 
by  the  ascultatory  method.  Woley  gives  the  average  systolic  pressure  as 
127  mm.  for  all  ages.  But  it  is  well  known  that  the  pressure  increases 
with  age  from  75  at  one  year,  to  105  in  youth  and  140  mm.  or  more  at  the 
age  of  fifty. 

AVERAGE  BLOOD  PRESSURE  MEASUREMENTS  or  MAN 


Standing 

Sitting 

Immediately 
after 
Exercise 

Systolic  Blood  Pressure: 

Men  

120  to  135 

no  to  133 

129  to  160 

Women  

105  to  127 

105  to  125 

118  to  150 

Diastolic  Blood  Pressure: 

Men 

68  to    90 

65  to    85 

70  to  i  20 

Women  

65  to     75 

60  to    80 

70  to    98 

The  Venous  Blood-Pressure  and  Capillary  Pressure. — The  blood- 
pressure  in  the  veins  is  nowhere  very  great,  but  is  greatest  in  the  small  veins, 
while  in  the  large  veins  near  the  heart  the  pressure  may  become  negative. 
In  other  words,  when  a  vein  is  put  in  connection  with  a  mercurial  manom- 
eter the  mercury  may  fall  in  the  arm  farthest  away  from  the  vein  and  will 
rise  in  the  arm  nearest  the  vein,  the  action  being  that  of  suction  rather  than 
pressure.  In  the  large  veins  of  the  neck  the  tendency  to  suck  in  air  is  espe- 
cially marked,  and  is  the  cause  of  death  in  some  accidents  or  operations 
in  that  region.  The  amount  of  pressure  in  the  brachial  vein  is  said  to 


228 


THE    CIRCULATION   OF   THE   BLOOD 


support  9  mm.  of  mercury,  whereas  the  pressure  in  the  veins  of  the  neck 
may  fall  to  a  negative  pressure  of  from  —3  to  —8  mm. 

The  variations  of  venous  pressure  during  systole  and  diastole  of  the 
heart  are  very  slight,  and  a  distinct  pulse  is  never  seen  in  veins  except  under 
extraordinary  circumstances.  In  certain  forms  of  cardiac  valvular  insuffi- 
ciency there  may  be  considerable  regurgitation  of  the  blood  with  a  strong 
venous  pulse. 

Careful  observations  upon  the  web  of  the  frog's  foot,  the  tongue  and 
mesentery  of  the  frog,  the  tails  of  newts  and  small  fishes,  and  upon  the 
skin  of  the  finger  behind  the  nail  (Hooker) ;  as  well  as  estimations  of  the 
amount  of  pressure  required  to  empty  the  vessels  of  blood  under  various 
conditions,  all  indicate  that  the  capillary  blood- pressure  is  subject  to  very 
great  variations.  Apparently  the  variations  follow  the  variations  of 
pressure  in  the  arteries,  though  the  measurements  of  the  capillary  pressure 
of  the  skin  in  man  indicate  that  it  is  occasionally  markedly  influenced  by 
the  venous  pressure  variations  (Hough).  In  the  skin  in  man  it  is  from  30 
to  50  mm.  mercury. 

The  pulse  in  the  arterioles,  capillaries,  and  venules  becomes  more  and 
more  evident  as  the  extravascular  pressure  is  increased.  The  pressure  in 
the  web  of  the  frog's  foot  has  been  found  to  be  equal  to  about  14  to  20  mm. 
of  mercury;  in  other  capillary  regions  the  pressure  is  found  to  be  equal  to 
from  one-fifth  to  one-half  of  the  ordinary  arterial  pressure. 


FIG.  191. — Schema  Showing  the  Relation  between  Blood  Pressure,  Velocity  of  Flow, 
and  Vascular  Area,  in  the  Arteries,  Capillaries,  and  Veins.  Ordinates  represent  height 
of  pressure  and  speed  of  flow.  The  abscissa,  b-c,  represents  zero  pressure  and  speed. 
Space  between  lines  a-b  and  d-e  represents  arterial  system;  between  d-e  and/-g,  capillary 
system,  and  between  f-g  and  h-i,  the  venous  system.  Line  A-B  equals  pressure;  line  C-D, 
speed  of  flow;  and  line  E-F,  vascular  area.  (Modified  from  Gad.) 

General  Variations  in  Blood-Pressure. — The  arterial  blood-pressure 
may  be  made  to  vary  by  alterations  in  either  of  the  chief  factors  upon  which 


THE    ARTERIAL    FLOW  2 29 

the  pressure  in  the  vessels  depends,  but  primarily  by  the  cardiac  contrac- 
tions and  the  peripheral  resistance.  Thus,  increase  of  blood-pressure  may 
be  brought  about  by  either,  i,  a  more  frequent  or  more  forcible  action  of 
the  heart,  or  2,  by  an  increase  of  the  peripheral  resistance.  On  the  other 
hand,  diminution  of  the  blood-pressure  may  be  produced,  either  by,  a,  a 
diminished  force  or  frequency  of  the  contractions  of  the  heart,  or  by  b,  a 
diminished  peripheral  resistance.  These  different  factors,  however,  al- 
though varying  constantly,  are  so  combined  that  the  general  arterial  pressure 
remains  fairly  constant.  For  example,  the  heart  may,  by  increased  force  or 
frequency  of  its  contractions,  distinctly  increase  the  blood  pressure,  but  this 
increased  action  is  almost  certainly  followed  by  diminished  peripheral  re- 
sistance, and  thus  the  two  altered  conditions  may  balance,  with  the  result 
of  bringing  back  the  blood-pressure  to  what  it  was  before  the  heart  began 
to  beat  more  rapidly  or  more  forcibly. 

It  will  be  clearly  seen  that  the  circulation  of  the  blood  within  the  blood- 
vessels must  depend  upon  the  diminution  of  the  pressure  from  the  heart 
to  the  capillaries,  and  from  the  capillaries  to  the  veins,  the  blood  flowing  in 
the  direction  of  least  resistance.  We  shall  presently  see  further  that  the 
local  flow  also  depends  upon  the  relations  between  the  heart's  action  and 
the  peripheral  resistance  both  general  and  local. 

The  Arterial  Flow. — The  character  of  the  flow  of  blood  through  the 
arterial  system  depends  to  a  very  considerable  extent  upon  the  structure 
of  the  arterial  walls,  and  particularly  upon  the  elastic  tissue  which  is  so  highly 
developed  in  them. 

The  elastic  tissue  of  the  arteries,  first  of  all,  guards  them  from  the  sud- 
denly exerted  pressure  to  which  they  are  subjected  at  each  contraction  of  the 
ventricles.  In  every  such  contraction,  as  is  above  seen,  the  contents  of  the 
ventricles  are  forced  into  the  arteries  more  quickly  than  they  are  discharged 
through  the  capillaries.  The  blood,  therefore,  being  for  an  instant  resisted 
in  its  onward  course,  a  part  of  the  force  with  which  it  is  impelled  is  directed 
against  the  sides  of  the  arteries;  under  this  force  their  elastic  walls  dilate, 
stretching  enough  to  receive  the  blood,  and  becoming  more  tense  and  more 
resisting  as  they  stretch.  Thus  by  yielding  they  break  the  shock  of  the 
force  impelling  the  blood.  On  the  subsidence  of  the  pressure,  should  the 
ventricles  cease  contracting,  the  arteries  are  able  by  the  same  elasticity  to 
resume  their  former  caliber. 

The  elastic  tissue  in  the  same  way  equalizes  the  current  of  blood  by  main- 
taining pressure  on  it  in  the  arteries  during  the  period  at  which  the  ventri- 
cles are  at  rest  or  are  dilating.  If  the  arteries  were  rigid  tubes,  the  blood, 
instead  of  flowing  as  it  does  in  a  constant  stream,  would  be  propelled  through 
the  arterial  system  in  a  series  of  spurts  corresponding  in  time  to  the  ventric- 
ular contractions  and  with  intervals  of  almost  complete  rest  during  the  in- 
action of  the  ventricles.  But  in  the  actual  condition  of  the  vessels,  the  force 
of  the  successive  contractions  of  the  ventricles  is  expended  partly  in  the 


230  THE    CIRCULATION    OF   THE   BLOOD 

direct  propulsion  of  the  blood  and  partly  in  the  dilatation  of  the  elastic  ar- 
teries; and  in  the  intervals  between  the  contractions  of  the  ventricles,  the 
force  of  the  recoil  is  employed  in  continuing  the  flow  onward.  Of  course 
the  pressure  exercised  is  equally  diffused  in  every  direction,  and  the  blood 
tends  to  move  backward  as  well  as  onward.  All  movement  backward, 
however,  is  prevented  by  the  closure  of  the  semilunar  valves,  which  takes 
place  at  the  very  commencement  of  the  recoil  of  the  arterial  walls. 

The  Arterial  Flow  is  Rhythmic. — By  the  exercise  of  the  elasticity 
of  the  arteries,  all  the  force  of  the  ventricles  is  expended  upon  the  circulation. 
That  part  of  the  force  which  is  used  up  or  rendered  potential  in  dilating  the 
arteries  is  restored  or  made  active  or  kinetic  when  they  recoil.  There  is  no 
loss  of  force,  neither  is  there  any  gain;  for  the  elastic  walls  of  the  artery  can- 
not originate  any  force  for  the  propulsion  of  the  blood;  they  only  restore 
that  which  they  receive  from  the  ventricles. 

Since  the  ventricular  discharge  is  intermittent,  there  will  be  intermittent 
accessions  of  pressure,  and  therefore  the  flow  of  blood  in  the  arteries  will  be 
periodically  accelerated.  The  volume  of  blood  discharged  from  a  cut  artery 
increases  and  decreases  with  the  systole  and  diastole  of  the  ventricles,  or  with 
the  systolic  and  diastolic  pressures  of  the  arteries  themselves,  the  maximal 
speed  being  at  the  moment  of  maximal  systolic  pressure,  see  page  228. 

The  equalizing  influence  of  the  resistance  of  the  successive  arterial 
branches  reacts  so  that  at  length  the  intermittent  accelerations  produced  in 
the  arterial  flow  by  the  discharge  of  the  heart  cease  to  be  observable,  and 
the  jetting  stream  is  converted  into  the  continuous  and  even  movement  of 
the  blood  which  characterizes  the  flow  in  the  capillaries  and  veins. 
The  resistance  which  is  offered  to  the  flow  of  the  blood  stream  in  these 
vessels  is  a  necessary  agent  in  the  production  of  a  continuous  stream  of 
blood  in  the  smaller  arteries  and  capillaries.  Were  there  no  greater  ob- 
stacle to  the  escape  of  blood  from  the  larger  arteries  than  exists  to  its  en- 
trance into  them  from  the  heart,  the  stream  would  be  intermittent, 
notwithstanding  the  elasticity  of  the  walls  of  the  arteries. 

The  muscular  element  of  the  middle  coat  co-operates  with  the  elastic 
element  in  adapting  the  caliber  of  the  vessels  to  the  quantity  of  blood  which 
they  contain;  for  the  amount  of  fluid  in  the  blood-vessels  varies  quite  con- 
siderably even  from  hour  to  hour,  and  can  never  be  quite  constant.  Were 
the  elastic  tissue  only  present,  the  pressure  exercised  by  the  walls  of  the 
containing  vessels  on  the  contained  blood  would  be  sometimes  very  small  and 
sometimes  inordinately  great.  The  presence  of  a  muscular  element,  how- 
ever, provides  for  a  certain  uniformity  in  the  amount  of  pressure  exercised: 
the  muscles  are  in  constant  action  or  tone,  and  it  is  by  this  adaptive,  uniform, 
gentle  muscular  contraction  that  the  normal  tone  of  the  blood-vessels  is 
maintained.  Deficiency  of  this  tone  is  the  cause  of  the  soft  and  yielding 
arterial  pulse,  and  the  sluggish  blood  flow  through  the  arterioles. 


THE    CAPILLARY    FLOW 


23I 


Incidentally  it  may  be  mentioned  that  the  elastic  and  muscular  contrac- 
tion of  an  artery  may  also  be  regarded  as  fulfilling  a  natural  purpose  when, 
the  artery  being  cut,  the  sudden  contraction  at  first  limits,  and  then,  in  con- 
junction with  the  coagulating  blood,  completely  arrests,  the  flow  of  blood. 
It  is  only  in  consequence  of  such  contraction  and  coagulation  that  we  are 
free  from  danger  through  even  very  slight  wounds;  for  it  is  only  when  the 
artery  is  closed  that  the  processes  for  the  more  permanent  and  secure  pre- 
vention of  bleeding  are  established. 

The  Velocity  of  the  Arterial  Blood  Flow. — The  velocity  of  the  blood 
current  at  any  given  point  in  the  various  divisions  of  the  circulatory  system 
is  inversely  proportional  to  the  united  sectional  area  at  that  point.  If  the 
united  sectional  area  of  all  the  branches  of  a  vessel  were  always  the  same 
as  the  area  of  the  vessel  from  which  they  arise,  and  if  the  aggregate  sectional 
area  of  the  capillary  vessels  were  equal  to  that  of  the  aorta,  the  mean  rapidity 
of  the  blood's  motion  in  the  small  arteries  and  in  the  capillaries  would  be  the 
same  as  in  the  aorta.  If  a  similar  correspondence  of  capacity  existed  in  the 
veins  there  would  be  an  equal  correspondence  in  the  rapidity  of  the  circula- 
tion in  them.  But  the  volume  of  the  arterial  and  venous  systems  may  be 
represented  by  two  truncated  cones  with  their  apices  directed  toward  the 
heart;  the  area  of  their  united  bases,  the  sectional  area  of  the  capillaries, 
being  about  eight  hundred  times  as  great  as  that  of  the  truncated  apex  rep- 
resenting the  aorta.  Thus  the  velocity  of  blood  in  the  smallest  arterioles 
and  the  capillaries  is  about  one-eight-hundredth  of  that  in  the  aorta. 

The  velocity  of  the  stream  of  blood  is  greatest  in  the  neighborhood  of 
the  heart.  The  rate  of  movement  is  greatest  during  the  ventricular  systole 
and  diminishes  during  the  diastole.  The  rate  of  flow  also  decreases  along 
the  arterial  system,  becoming  least  in  the  parts  of  the  system  most  distant 
from  the  heart.  Chauveau  has  estimated  the  rapidity  of  the  blood  stream 
in  the  carotid  of  the  horse  at  over  20  inches  per  second  during  the  heart's 
systole,  and  nearly  6  inches  during  the  diastole  (520-150  mm.)  see  figure  191. 

The  Capillary  Flow. — It  is  in  the  capillaries  that  the  chief  resistance 
is  offered  to  the  progress  of  the  blood;  for  in  them  the  friction  of  the  blood 
is  greatly  increased  by  the  enormous  multiplication  of  the  surface  with  which 
it  is  brought  in  contact. 

When  the  capillary  circulation  is  examined  in  any  transparent  part  of  a 
full-grown  living  animal  by  means  of  the  microscope,  figures  193,  194,  the 
blood  is  seen  to  flow  with  a  constant  equable  motion ;  the  red  blood  corpus- 
cles moving  along,  mostly  in  single  file,  and  bending  in  various  ways  to  ac- 
commodate themselves  to  the  tortuous  course  of  the  capillary,  but 
instantly  recovering  their  normal  outline  on  reaching  a  wider  vessel. 

At  the  circumference  of  the  stream  and  adhering  to  the  walls  of  the 
larger  capillaries,  but  especially  well  marked  in  the  small  arteries  and  veins, 
there  is  a  layer  of  plasma  which  appears  to  be  motionless.  The  existence 


232  THE    CIRCULATION   OF   THE    BLOOD 

of  this  still  layer,  as  it  is  termed,  is  inferred  both  from  the  general  fact  that 
such  a  one  exists  in  all  fine  tubes  traversed  by  fluid,  and  from  what  can  be 
seen  in  watching  the  movements  of  the  blood  corpuscles.  The  red  cor- 
puscles occupy  the  middle  of  the  stream  and  move  with  comparative 
rapidity;  the  colorless  corpuscles  run  much  more  slowly  by  the  walls  of  the 
vessels;  while  next  to  the  wall  there  is  a  transparent  space  in  which  the 
fluid  appears  to  be  at  rest ;  for  if  any  of  the  corpuscles  happen  to  be  forced 
within  it,  they  move  more  slowly  than  before,  rolling  lazily  along  the  side 
of  the  vessel  and  often  adhering  to  its  wall,  figure  194.  Part  of  this  slow 
movement  of  the  colorless  corpuscles  and  their  occasional  stoppage  may  be 


4-fFfWKr 


FIG.  193. — Capillary  Network  from  Human  Pia  Mater,  Showing  also  an  Arteriole  in 
Optical  Section";  and  a  Small  Vein.      X  350.     A,  Vein;  B,  arteriole;  C,  large  capillary 
D,  small  capillaries.     (Bailey.) 

due  to  their  having  a  tendency  to  adhere  to  the  walls  of  the  vessels.  Some- 
times, indeed,  when  the  motion  of  the  blood  is  not  strong,  many  of  the 
white  corpuscles  collect  in  a  capillary  vessel,  and  for  a  time  entirely 
prevent  the  passage  of  the  red  corpuscles. 

When  the  peripheral  resistance  is  greatly  diminished  by  the  dilatation 
of  the  small  arteries  and  capillaries,  so  much  blood  passes  on  from  the 
arteries  into  the  capillaries  at  each  stroke  of  the  heart  that  there  is  not 
sufficient  remaining  in  the  arteries  to  distend  them.  Thus,  the  intermit- 
tent current  of  the  ventricular  systole  is  not  always  converted  into  a  con- 
tinuous stream  by  the  elasticity  of  the  arteries  before  the  capillaries  are 
reached.  The  intermittency  of  the  flow  occurs  both  in  capillaries  and 
veins  and  a  venous  pulse  is  produced.  The  same  phenomenon  may  occur 
when  the  arteries  become  rigid  from  disease,  and  when  the  beat  of  the 


THE    CAPILLARY    FLOW 


233 


heart  is  so  slow  or  so  feeble  that  the  blood  at  each  cardiac  systole  has  time 
to  pass  on  to  the  capillaries  before  the  next  stroke  occurs.  The  amount  of 
blood  sent  at  each  stroke  is  not  sufficient  properly  to  distend  the  elastic 
arteries. 

It  was  formerly  supposed  that  the  occurrence  of  any  transudation 
from  the  interior  of  the  capillaries  into  the  midst  of  the  surrounding  tissues 
was  confined,  in  the  absence  of  injury,  strictly  to 
the  fluid  part  of  the  blood;  in  other  words,  that 
the  corpuscles  could  not  escape  from  the  circulating 
stream,  unless  the  wall  of  the  containing  blood  ves- 
sel was  ruptured.  It  is  true  that  the  English 
physiologist  Augustus  Waller  affirmed  in  1846  that 
he  had  seen  blood  corpuscles,  both  red  and  white, 
pass  bodily  through  the  wall  of  the  capillary  vessel 
in  which  they  were  contained  (thus  confirming  what 
had  been  stated  a  short  time  previously  by  Addi- 
son).  He  said  that  no  opening  could  be  seen 
before  their  escape  and  that  none  could  be  observed 
afterward,  so  rapidly  was  the  part  healed.  But 
these  observations  did  not  attract  much  notice 
until  the  phenomenon  of  escape  of  the  blood  cor- 
puscles from  the  capillaries  and  minute  veins,  apart 
from  mechanical  injury,  was  rediscovered  by  Cohn- 
heim  in  1867. 

Cohnheim's  experiment  demonstrating  the  pas- 
sage of  the  corpuscles  through  the  wall  of  the  blood 
vessel  is  performed  in  the  following  manner:  A 
frog  is  curarized;  that  is  to  say,  paralysis  is  pro- 
duced by  injecting  under  the  skin  a  minute  quantity  of  the  poison  called 
curare.  The  abdomen  is  then  opened,  a  portion  of  the  small  intestine  is 
drawn  out,  and  its  transparent  mesentery  spread  out  under  a  microscope. 
After  a  variable  time,  occupied  by  dilatation  following  contraction  of  the 
minute  vessels  and  the  accompanying  quickening  of  the  blood  stream, 
there  ensues  a  retardation  of  the  current  and  the  red  and  white  blood 
corpuscles  begin  to  make  their  way  through  the  capillaries  and  small  veins. 

The  white  corpuscles  pass  through  the  capillary  wall  chiefly  by  the 
ameboid  movement  with  which  they  are  endowed.  This  migration 
occurs  to  a  limited  extent  in  health,  but  in  inflammatory  conditions  is 
much  increased. 

The  process  of  diapedesis  of  the  red  corpuscles,  which  occurs  under  cir- 
cumstances of  impeded  venous  circulation,  and  consequently  increased 
blood  pressure,  resembles  closely  the  migration  of  the  leucocytes,  with  the 
exception  that  they  are  squeezed  through  the  wall  of  the  vessel,  and  do  not, 
like  the  leucocytes,  work  their  way  through  by  ameboid  movement. 


FIG.  194. — A  Large 
Capillaryfrom  the  Frog's 
Mesentery  Eight  Hours 
after  Irritation  had  been 
set  up,  Showing  Emi- 
gration of  Leucocytes. 
a,  Cells  in  the  act  of 
traversing  the  capillary 
wall;  b,  some  already 
escaped.  (Frey.) 


234  THE    CIRCULATION   OF   THE   BLOOD 

Various  explanations  of  these  remarkable  phenomena  have  been 
suggested.  It  is  no  longer  believed  that  pseudo-stomata  between  contigu- 
ous endothelial  cells  provide  the  means  of  escape  for  the  blood  corpuscles. 
The  chief  share  in  the  process  is  probably  due  to  mobility  and  con- 
traction of  the  parts  concerned,  both  of  the  corpuscles  and  of  the  capillary 
wall  itself. 

The  Speed  of  the  Blood  in  the  Capillaries. — The  velocity  of  the  blood 
through  the  capillaries  must,  of  necessity,  be  largely  influenced  by  that 
which  occurs  in  the  vessels  on  both  sides  of  them,  in  the  arteries  and 
the  veins.  Their  intermediate  position  causes  them  to  respond  at  once  to 
any  alteration  in  the  size  or  rate  of  the  arterial  or  venous  blood  stream. 
Thus,  the  apparent  contraction  of  the  capillaries,  on  the  application  of 
certain  irritating  substances  or  during  certain  mental  states,  and  their 
dilatation  in  blushing  may  be  referred  primarily  to  the  corresponding 
action  of  the  small  arteries. 

The  Measurement  of  Velocity  in  the  Capillaries. — The  observation  of 
Hales,  E.  H.  Weber,  and  Valentin  agree  very  closely  as  to  the  rate  of  the 
blood  current  in  the  capillaries  of  the  frog.  The  mean  of  their  estimates 
gives  the  velocity  of  the  systemic  capillary  circulation  at  about  0.5  mm.  per 
second.  The  velocity  in  the  capillaries  of  warm-blooded  animals  is 
greater,  in  the  dog  0.5  to  0.75  mm.  per  second.  This  may  seem  incon- 
sistent with  the  facts,  which  show  that  the  whole  circulation  is  accom- 
plished in  about  half  a  minute.  But  the  whole  length  of  capillary  vessels, 
through  which  any  given  portion  of  blood  has  to  pass,  probably  does  not 
exceed  0.5  mm.  Therefore  the  time  required  for  each  quantity  of  blood  to 
traverse  its  own  appointed  portion  of  the  general  capillary  system  will 
scarcely  amount  to  more  than  a  second.  This  comparatively  slow 
velocity  is  evidently  favorable  to  the  nutritive  interchanges  that  go  on 
through  these  thin-walled  vessels  between  the  blood  within  the  capillaries 
and  the  outside  active  tissues. 

The  Venous  Flow. — The  blood  current  in  the  veins  is  maintained, 
a,  primarily  by  the  contractions  of  the  left  ventricle;  but  very  effectual 
assistance  to  the  flow  is  afforded,  &,  by  the  action  of  the  muscles  capable  of 
pressing  on  the  veins  with  valves,  and  c,  by  the  aspiration  of  the  thorax 
and  possibly,  d,  by  the  aspiration  of  the  heart  itself. 

The  effect  of  muscular  pressure  upon  the  circulation  may  be  thus  ex- 
plained: When  pressure  is  applied  to  any  part  of  a  vein,  and  the  current  of 
blood  in  it  is  obstructed,  the  portion  behind  the  seat  of  pressure  becomes 
swollen  and  distended  as  far  back  as  the  next  pair  of  valves,  which  are  in 
consequence  closed.  Thus,  whatever  force  is  exercised  by  the  external 
pressure  of  the  muscles  on  the  veins,  is  distributed  partly  in  pressing  the 
blood  onward  in  the  proper  course  of  the  circulation,  and  partly  in  pressing 
it  backward  and  closing  the  valves  behind. 


THE   VELOCITY    OF    THE    CIRCULATION  235 

The  circulation  might  lose  as  much  as  it  gains  by  such  an  action  if  it 
were  not  for  the  numerous  communications,  or  venous  anastomoses. 
Owing  to  these  anastomoses  the  closing  up  of  the  venous  channel  by  the 
backward  pressure  is  prevented  from  being  any  serious  hindrance  to  the 
circulation,  since  the  blood  which  is  arrested  in  its  onward  course  by  the 
closed  valves  can  at  once  pass  through  some  anastomosing  channel  and 
proceed  on  its  way  by  another  vein.  Thus  the  effect  of  muscular  pressure 
upon  veins  which  have  valves  is  turned  almost  entirely  to  the  advantage 
of  the  circulation.  The  pressure  of  the  blood  onward  is  all  advantageous, 
and  the  pressure  of  the  blood  backward  is  prevented  from  being  a  hin- 
drance by  the  closure  of  the  valves  and  by  the  anastomoses  of  the  veins. 

The  venous  flow  is  also  assisted  by  the  aspiration  of  the  thorax  and  to 
some  extent  by  that  of  the  heart,  since  at  some  time  during  every  cardiac 
cycle  the  intra-auricular  and  intra-ventricular  pressure  falls  below  that  of 
the  atmosphere.  This  activity  will  be  considered  more  fully  in  the  chapter 
on  Respiration.  In  this  connection  it  may  be  said,  however,  that  the  pres- 
sure in  the  great  veins  falls  during  inspiration  and  rises  during  expiration. 

The  Velocity  in  the  Veins. — The  velocity  of  the  blood  is  greater  in  the 
veins  than  in  the  capillaries,  but  less  than  in  the  arteries;  this  fact  depend- 
ing upon  the  relative  capacities  of  the  arterial  and  venous  systems.  If 
an  accurate  estimate  of  the  proportionate  areas  of  arteries  and  the  veins 
corresponding  to  them  could  be  made,  we  might,  from  the  velocity  of  the 
arterial  current,  calculate  that  of  the  venous.  The  usual  estimation  is  that 
the  capacity  of  the  veins  is  about  two  or  three  times  as  great  as  that  of  the 
arteries,  and  that  the  velocity  of  the  blood's  motion  is,  therefore,  about 
one-half  or  one-third  as  great  in  the  veins  as  in  the  arteries,  i.e.,  200  mm. 
a  second.  The  rate  at  which  the  blood  moves  in  the  smallest  venules  is 
only  slightly  greater  than  that  in  the  capillaries,  but  the  speed  of  flow 
gradually  increases  the  nearer  the  vessel  approaches  to  the  heart.  The 
total  sectional  area  of  the  venous  trunks,  compared  with  that  of  the 
branches  opening  into  them,  becomes  gradually  smaller  as  the  trunks 
advance  toward  the  heart,  figure  191. 

The  Velocity  of  the  Circulation  as  a  Whole. — It  would  appear  that 
a  portion  of  blood  can  traverse  the  entire  course  of  the  circulation,  in  the 
horse,  in  half  a  minute.  Of  course  it  would  require  longer  to  traverse 
the  vessels  of  the  most  distant  part  of  the  extremities  than  to  go  through 
those  of  the  neck,  but  taking  an  average  length  of  the  vessels  to  be 
traversed  it  may  be  concluded  that  half  a  minute  represents  the  average 
rate.  Stewart  estimated  that  the  circulation  time  in  man  is  probably  not 
less  than  twelve  nor  more  than  fifteen  seconds. 

Satisfactory  data  for  these  estimates  are  afforded  by  the  results  of. expe- 
riments to  ascertain  the  rapidity  with  which  chemicals  introduced  into  the 
blood  are  transmitted  from  one  part  of  the  vascular  system  to  another.  The 


236.  THE    CIRCULATION    OF   THE   BLOOD 

time  required  for  the  passage  of  solutions  of  potassium  ferrocyanide,  mixed 
with  the  blood,  from  one  jugular  vein,  through  the  right  side  of  the  heart,  the 
pulmonary  vessels,  the  left  cavities  of  the  heart,  and  the  general  circulation, 
to  the  jugular  vein  of  the  opposite  side,  varies  from  twenty  to  thirty  seconds 
in  the  dog.  The  same  substance  is  transmitted  from  the  jugular  vein  to  the 
great  saphenous  vein  in  twenty  seconds;  from  the  jugular  vein  to  the  mes- 
enteric  artery  in  between  fifteen  and  thirty  seconds;  to  the  facial  artery, 
in  one  experiment,  in  between  ten  and  fifteen  seconds;  in  another  experi- 
ment, in  between  twenty  and  twenty- five  seconds;  in  its  transit  from  the 
jugular  vein  to  the  metatarsal  artery,  it  occupies  between  twenty  and 
thirty  seconds.  The  result  is  said  to  be  nearly  the  same  whatever  the 
rate  of  the  heart's  action.  In  more  recent  methods  some  innocuous  dye 
like  methylene  blue  is  used,  since  it  permits  the  determination  without 
the  loss  of  blood,  the  change  in  color  being  visible  through  the  walls  of  the 
blood  vessels. 

Stewart  has  made  most  accurate  measurements  of  the  circulation  time 
by  the  electrical- resistance  method.  Strong  salt  solutions  injected  into  the 
jugular  vein  on  one  side  when  they  reach  the  other  jugular  (or  any  other 
vessel)  are  instantly  detected  by  a  decrease  in  the  electrical  resistance  through 
the  vessel  when  it  is  laid  between  the  poles  of  the  proper  conductivity 
apparatus. 

In  all  these  experiments  it  is  assumed  that  the  substance  injected  moves 
with  the  blood  and  at  the  same  rate,  and  does  not  move  from  one  part  of 
the  organs  of  circulation  to  another  by  diffusing  itself  through  the  blood  or 
tissues  more  quickly  than  the  blood  moves.  The  assumption  may  be  ac- 
cepted that  the  times  above  mentioned  as  occupied  in  the  passage  of  the  in- 
jected substances  are  the  times  in  which  the  portion  of  blood  itself  is  carried 
from  one  part  to  another  of  the  vascular  system. 

Another  mode  of  estimating  the  general  velocity  of  the  circulating  blood 
is  by  calculating  it  from  the  quantity  of  blood  supposed  to  be  contained  in 
the  body  and  from  the  quantity  which  can  pass  through  the  heart  in  each 
of  its  contractions.  But  the  conclusions  arrived  at  by  this  method  are  less 
satisfactory.  For  the  total  quantity  of  blood  and  the  capacity  of  the  cavities 
of  the  heart  have  as  yet  been  only  approximately  ascertained.  Still  the  most 
careful  of  the  estimates  thus  made  accord  very  nearly  with  those  already 
mentioned;  and  it  may  be  assumed  that  the  blood  may  all  pass  through 
the  heart  in  man  in  about  thirty  seconds  or  even  less. 

THE  PULSE. 

The  most  characteristic  feature  of  the  arterial  pressure  and  blood 
flow  is  its  intermittency,  and  this  intermittent  flow  is  seen  or  felt  as 
waves  of  change  in  diameter  of  the  arteries,  known  as  the  Pulse. 


THE    SPHYGMOGRAPH 


237 


The  pulse  is  generally  described  as  a  wave-like  expansion  of  the  artery 
produced  by  the  injection  of  blood  at  each  ventricular  systole  into  the  already 
full  aorta.  The  force  of  the  left  ventricle  is  expended  in  pressing  the  blood 
forward  and  in  dilating  the  aorta.  With  the  injection  of  each  new  quantity 
of  blood  into  the  aorta  there  is  a  wave  of  dilatation  which  passes  on,  expand- 
ing the  arteries  as  it  goes,  running  as,  it  were,  over  the  more  slowly  traveling 
blood  contained  in  them,  and  producing  the  pulse  as  it  proceeds.  A  sharp 
distinction  must  be  made  between  the  passage  of  the  pulse  wave  along  an 
artery  and  the  rate  of  flow  of  the  blood  in  the  vessel.  The  pulse  produced  by 
any  given  beat  of  the  heart  is  not  felt  at  the  same  moment  in  all  parts  of  the 
body.  Thus,  it  can  be  felt  in  the  carotid  a  short  time  before  it  is  perceptible 
in  the  radial  artery,  and  in  this  vessel  before  it  occurs  in  the  dorsal  artery  of 
the  foot.  Careful  measurements  of  the  intervals  between  the  time  of  the 
pulse  at  the  carotid  and  at  the  wrist  shows  that  the  delay  in  the  beat  is  in 
proportion  to  the  distance  of  the  artery  from  the  heart.  The  difference  in 
time  between  the  pulse  of  any  two  arteries  probably  never  exceeds  one-sixth  to 
one-eighth  of  a  second.  The  rate  at  which  the  pulse  travels  in  the  arteries 
is  from  five  to  ten  meters  per  second. 

The  distention  of  each  artery  increases  both  its  length  and  its  diameter. 
In  their  elongation  the  arteries  change  their  form,  the  straight  ones  becoming 
slightly  curved,  and  those  already  curved  becoming  more  so;  but  they  re- 
cover their  previous  form  as  well  as  their  diameter  when  the  ventricular 
contraction  ceases,  and  their  elastic  walls  recoil.  The  increase  of  their 
curves  which  accompanies  the  distention  of  arteries,  and  the  succeeding 
recoil,  may  be  well  seen  in  the  prominent  temporal  artery  of  an  old  person. 
In  feeling  the  pulse,  the  finger  cannot  distinguish  the  sensation  produced 
by  the  dilatation  from  that  produced  by  the  elongation  and  curving.  That 
which  it  perceives  most  plainly,  however,  is  the  dilatation  and  return  more 
or  less  to  the  cylindrical  form,  of  the  artery  which  has  been  partially  flattened 
by  the  finger. 

The  Sphygmograph. — Much  light  has  been  thrown  on  what  may 
be  called  the  form  of  the  pulse  wave  by  an  instrument  called  the  sphygmo- 
graph,  figures  195  and  196.  The  principle  on  which  it  acts  will  be  seen 
on  reference  to  the  figures. 

A  small  button  replaces  the  finger  in  the  act  of  taking  the  pulse.  This 
button  is  made  to  rest  lightly  on  the  artery  the  pulsations  of  which  it  is  de- 
sired to  investigate.  The  up-and-down  movement  of  the  button  is  com- 
municated to  the  lever,  to  the  hinder  end  of  which  is  attached  a  light  spring. 
The  spring  is  adjusted  to  the  proper  tension  to  follow  the  movements  of  the 
artery  wall  during  the  pulse  wave.  The  sphygmograph  is  bound  on  the 
wrist  while  taking  a  record. 

It  is  evident  that  the  beating  of  the  pulse  will  cause  an  up-and-down 
movement  of  the  lever,  the  pen  of  which  will  write  the  effect  on  a  smoked 


238 


THE    CIRCULATION    OF    THE   BLOOD 


card  moved  by  the  clock-work  of  the  instrument. 

Thus  a  tracing  of  the  pulse  is  obtained,  and  in  this  way  much  more  deli- 
cate changes  can  be  seen  than  can  be  felt  by  the  mere  application  of  the  finger. 


FIG.  195. — Diagram  of  the  Lever  of  the  Sphygmograph. 

The  principle  of  the  sphygmometer  of  Roy  and  Adami  is  shown  in  the  diagram,  figure 
197. 

The  apparatus  consists  of  a  box,  a,  which  is  moulded  to  fit  over  the  end  of  the  radius 
so  as  to  bridge  over  the  radial  artery.  Within  this  is  a  flexible  bag,  q,  filled  with  water, 


FIG.  196. — Dudgeon's  Sphygmograph. 

and  connected  by  a  T-tube  with  a  rubber  bag,  h,  and  mercurial  manometer.  The  fluid 
in  the  box  may  be  raised  to  any  desired  pressure,  and  may  then  be  shut  off  by  tap,  c. 
At  the  upper  part  of  the  box  is  a  circular  opening,  and  resting  upon  b  is  a  flat  button,  d, 
which  by  means  of  a  short  light  rod,  e,  communicates  the  movement  of  b  to  the  lever,/. 
At  the  axis  of  rotation  of  this  lever  is  a  spiral  watch-spring,  g,  which  can  be  tightened 
at  will,  so  that  the  lever  can  be  made  to  take  a  vertical  position  at  any  desired  hydro- 
static pressure  within  the  box.  The  movements  of  the  lever  are  recorded  upon  a  piece 
of  blackened  glazed  paper  made  to  move  in  a  vertical  direction  past  it.  When  in  use, 
the  box  is  fixed  upon  the  wrist  by  an  appropriate  holder,  and  the  pressure  is  raised  to 


SPHYGMOGRAM 


239 


any  desired  height  to  which  the  lever  is  adapted  by  tightening  or  slackening  the  spring ; 
the  tap,  c,  is  then  closed.  The  pressure  within  the  box  acts  in  all  directions,  and  is 
correctly  indicated  by  the  manometer. 


To  manometer. 


FIG.  197.— Diagrammatic  Sectional  Representation  of  the  Sphygmometer.  a,  Box 
by  which  the  portion  of  the  artery  is  covered;  &,  thin- walled  india-rubber  bag  filled  with 
water,  and  communicating  through  tap,  c,  with  the  manometer  and  thick- walled  rubber  bag, 
h;  d,  piston  connected  by  rod,  e,  with  recording  lever,/;  g,  spiral  spring,  attached  to  axis  of 
lever,  and  by  which  the  pressure  in  b,  against  the  piston,  d,  is  counterbalanced;  k,  skin  and 
subcutaneous  tissue;  m,  end  of  radius  seen  in  section;  n,  radial  artery  seen  in  section. 
(Roy  and  Adami.) 


Sphygrnogram. — The  tracing  of  the  pulse  obtained  by  the  use  of  the 
sphygmograph,  called  a  sphymogram,  differs  somewhat  according  to  the  artery 
from  which  it  is  taken,  but  its  general  characters  are  much  the  same  in  all 
cases.  It  consists  of  a  sudden  upstroke,  or  anacrotic  limb,  figure  198,  A, 
which  is  somewhat  higher  and  more  abrupt  in  the  pulse  of  the  carotid  and  of 
other  arteries  near  the  heart  than  in  the  radial  and  other  arteries  more  re- 
mote; and  a  gradual  decline  or  catacrotic  limb,  P,  less  abrupt,  and  taking  a 
longer  time  than  A.  It  is  seldom,  however,  that  the  decline  is  an  uninter- 
rupted fall;  it  is  usually  marked  about  half-way  by  a  distinct  notch,  C-D 
the  dicrotic  notch,  followed  immediately  by  a  second  more  or  less  marked 
ascent  of  the  lever  called  the  dicrotic  wave,  D.  Not  infrequently  there  is 
also  at  the  beginning  of  the  descent  a  slight  wave  previous  to  the  dicrotic 
notch;  this  is  called  the  pre-dicrotic  wave,  and  in  addition  there  may  be  one 
or  more  slight  waves  after  the  dicrotic,  called  post-dicrotic,  E.  The  inter- 
ruptions in  the  downstroke  are  called  the  catacrotic  waves  to  distinguish 
them  from  an  interruption  in  the  upstroke,  called  the  anacrotic  wave,  which 
is  sometimes  met  with. 


240  THE    CIRCULATION   OF   THE   BLOOD 

The  explanation  of  these  tracings  present  some  difficulties,  not,  how- 
ever, as  regards  the  two  primary  factors,  viz.,  the  upstroke  and  downstroke, 


FIG.  198.— Diagram  of  Pulse  Tracing.     A,  upstroke  or  anacrotic  limb;  P,  downstroke  or 
katacrotic  limb;  C,  pre-dicrotic  wave;  D,  dicrotic;  E,  post-dicrotic  wave. 

because  they  are  universally  taken  to  mean  the  sudden  injection  of  blood 
into  the  already  distended  arteries,  and  the  gradual  recovery  of  the  arteries 
by  their  recoil.  These  points  may  be  demonstrated  on  a  system  of  elastic 
tubes,  with  a  pump  to  inject  water  at  regular  intervals,  just  as  well  as  on  the 
radial  artery,  or  on  the  arterial  schema,  a  more  complicated  system  of  tubes 
in  which  the  heart,  the  arteries,  the  capillaries,  and  veins  are  represented. 
If  we  place  two  or  more  sphygmographs  upon  such  a  system  of  tubes  at  in- 
creasing distances  from  the  pump,  we  may  demonstrate,  first,  that  the  rise 


FIG.  199. — Sphygmogram  from  the  Radial  Artery  Taken  with  Marey's  Sphygmograph. 

(Langendorff.) 

of  the  lever  commences  earliest  in  that  nearest  the  pump,  and,  second,  that 
it  is  higher  and  more  sudden.  So  in  the  arteries  of  the  body  the  wave  gradu- 
ally gets  less  and  less  as  we  approach  the  periphery  of  the  arterial  system, 
and  it  is  lost  in  the  capillaries. 

The  origin  of  the  secondary  waves  is  to  some  extent  a  matter  of  uncer- 
tainty. The  anacrotic  wave  occurs  when  the  peripheral  resistance  is  high; 
that  is,  when,  for  some  time  during  the  systole,  the  flow  from  the  aorta  toward 
the  periphery  is  slower  than  the  flow  from  the  ventricle  into  the  aorta.  Thus 
it  is  seen  in  some  cases  of  nephritis  where  the  arteries  are  rigid  and  the  periph- 
eral resistance  is  high. 

The  dicrotic  wave  is  the  most  important  of  the  secondary  waves,  and 
has  been  the  subject  of  much  discussion.  It  is  constantly  present  in  pulse 
tracings,  but  varies  in  height.  In  point  of  time  the  dicrotic  wave  occurs 
immediately  after  the  closure  of  the  aortic  semilunar  valves.  In  certain 


SPHYGMOGRAM 


24I 


conditions,  generally  of  disease,  it  becomes  so  marked  as  to  be  quite  plain 
to  the  unaided  finger.  Such  a  pulse  is  called  dicrotic.  The  generally  ac- 
cepted explanation  of  the  cause  of  the  dicrotic  wave  is  that  it  represents  a 


B 


FIG.  200. — A ,  Normal  Pulse  Tracing  from  Radial  of  Healthy  Adult  Obtained  by  the 
Sphygmometer;  B,  from  same  artery,  with  the  same  extra-arterial  pressure,  taken  during 
acute  nasal  catarrh.  1-2  Anacrotic  limb;  2-8  Catacrotic  limb;  3  Predicrotic  notch; 
5  Dicrotic  crest;  6  Postdicrotic  notch;  7  Postcrotic  crest;  4  Dicrotic  notch. 

rebound  of  the  overdistended  artery  at  the  time  of  the  closure  of  the  aortic 
valves.  During  systole,  as  the  blood  is  forcibly  injected  into  the  aorta, 
there  is  an  overdistention  of  the  artery.  The  systole  suddenly  ends,  the 
aorta  by  reason  of  its  elasticity  tends  to  recover  itself,  the  blood  is  driven 
back  against  the  semilunar  valves,  closing  them  and  at  the  same  time  giv- 
ing rise  to  a  wave,  the  dicrotic  wave,  which  begins  at  the  heart  and  travels 
onward  toward  the  periphery  like  the  primary  wave.  According  to  Foster, 
the  conditions  favoring  the  development  of  dicrotism  are:  i,  a  highly  ex- 
tensible and  elastic  arterial  wall;  2,  a  comparatively  low  mean  blood-press- 
ure, leaving  the  extensible  reaction  free  scope  to  act;  3,  a  vigorous  and  rapid 
stroke  of  the  ventricle  discharging  into  the  aorta  a  considerable  quantity 
of  blood.  The  other  secondary  waves  are  probably  due  to  the  oscillations 
in  the  elastic  recoil  of  the  arteries,  though  some  of  them  at  least  may  be 
due  to  the  inertia  of  the  instruments  used. 

In  the  use  of  the  sphygmograph  care  must  be  taken  in  the  regulation  of 
the  pressure  of  the  spring.  If  the  pressure  be  too  great,  the  characters  of 
the  pulse  may  be  almost  entirely  obscured  or  the  artery  may  be  completely 
obstructed  and  no  tracing  is  obtained.  On  the  other  hand,  if  the  pres- 


242  THE    CIRCULATION    OF    THE    BLOOD 

sure  is  too  slight,  a  very  small  part  of  the  characters  may  be  represented  on 
the  tracing. 

THE   PERIPHERAL   REGULATION   OF   THE   FLOW   OF   BLOOD. 

The  flow  of  blood  through  the  circulatory  system  depends  on  the  inter- 
action of  several  factors  which  have  already  been  mentioned  in  another  con- 
nection: The  rate  and  volume  of  the  heart-beat,  the  elasticity  of  the  blood 
vessels,  the  resistance  of  the  microscopic  peripheral  vessels,  and  the  volume 
of  blood  in  the  body.  We  have  already  learned,  page  205,  that  both  the 
rate  and  volume  of  the  contractions  of  the  heart  are  under  very  minute 
and  intimate  regulation  and  control  through  the  cardiac  nervous  mechanism. 
Also  we  have  found  that  there  is  intimate  co-ordination  between  the  activity 
of  the  heart  and  the  activity  of  all  other  parts  of  the  body,  a  co-ordination 
secured  through  the  nervous  system.  All  regulation  which  affects  the 
heart  must  of  necessity  affect  the  general  blood-pressure  and,  therefore, 
not  directly  any  particular  part. 

The  general  elasticity  of  the  blood  vessels,  and  of  the  arteries  in  par- 
ticular, which  makes  the  general  arterial  pressure  possible,  is  dependent 
primarily  on  the  presence  of  a  large  amount  of  elastic  connective  tissue  in 
the  walls  of  the  vessels.  The  elasticity  of  this  tissue  is  a  purely  passive 
property  which  can  be  utilized  only  by  some  positive  source  of  energy,  in 
this  instance  the  heart. 

The  Variations  in  Peripheral  Resistance.  —  Certain  arteries  and 
veins,  especially  the  smallest  ones,  the  arterioles  and  venules,  are  supplied 
with  muscular  tissue  in  their  walls.  The  activity  of  these  muscles  in  the 
vascular  complex  makes  the  peripheral  regulation  of  the  flow  of  blood 
possible.  This  muscular  tissue  not  only  exhibits  a  passive  elasticity 
comparable  to  that  of  the  yellow  elastic  connective  tissue,  but  upon  the 
proper  stimulation  it  actively  contracts  or  relaxes,  thus  securing  to  the  per- 
ipheral resistance  of  the  vessels  an  active  adjustment  to  the  ever- varying 
dynamic  conditions  of  the  vascular  apparatus. 

The  muscular  tissue  in  the  vessels  increases  relatively  in  amount  as 
the  vessels  become  smaller.  In  the  arterioles  it  is  developed  out  of  all 
proportion  to  the  other  elements.  In  fact,  in  passing  from  the  arterioles 
to  the  capillary  vessels,  made  up  as  we  have  seen  of  endothelial  cells  with  a 
supporting  ground  substance  only,  the  last  change  on  the  side  of  the  arteries, 
which  occurs  as  the  vessels  become  smaller,  is  the  disappearance  of 
muscular  fibers. 

The  office  of  the  muscular  coat  is  to  adjust  the  size  of  the  arterioles  and, 
therefore,  the  flow  of  the  blood.  It  is  to  regulate  the  quantity  of  blood  to  be 
received  by  each  part  or  organ,  and  to  adjust  this  quantity  to  the  requirements 
of  each,  according  to  various  circumstances,  but  chiefly  according  to  the  de- 


THE    DISCOVERY    OF    THE   VASO-MOTOR    NERVES  243 

gree  of  activity  which  each  organ  at  different  times  exhibits.  The  amount  of 
work  done  by  each  organ  of  the  body  constantly  varies,  and  the  variations 
often  quickly  succeed  each  other,  so  that,  as  in  the  muscles  for  example, 
within  the  same  hour  a  part  may  be  now  very  active  and  now  quite  inactive. 
In  all  its  active  exercise  of  function,  such  an  organ  requires  a  larger  supply  of 
blood  than  is  sufficient  for  it  during  the  times  when  it  is  comparatively 
inactive. 

It  is  evident  that  the  heart  cannot  regulate  the  blood-supply  to  each 
part  of  the  body  at  different  periods  independently  of  the  other  parts. 
Neither  could  this  be  regulated  by  any  general  and  uniform  contraction  of 
the  arteries.  But  it  may  be  regulated  by  the  power  which  the  arteries  of 
each  part  have,  through  their  muscular  tissue,  of  contracting  or  relaxing 
so  as  to  diminish  or  increase  their  size.  Since  the  general  blood  pressure 
is  fairly  constant  the  size  of  the  local  vessels  controls  the  supply  of  blood  to 
the  particular  part  of  the  body  to  which  the  vessels  are  distributed.  Thus, 
while  the  ventricles  of  the  heart  determine  the  total  quantity  of  blood  to  be 
sent  onward  to  each  contraction  and  the  force  of  its  propulsion,  and  while 
the  large  and  merely  elastic  arteries  distribute  the  blood  and  equalize  its 
stream,  the  smaller  arteries  regulate  and  determine  the  .proportion  of  the 
whole  quantity  of  blood  which  shall  be  distributed  to  each  particular 
organ. 

The  variation  of  the  size  of  arterioles  and,  therefore,  of  the  resistance 
to  the  flow  of  the  blood  in  them  is  secured  by  the  contractions  of  the  mus- 
cular tissue,  but  the  muscles  are  regulated  in  their  contractions  by  the 
nervous  system.  The  muscular  tissue  in  the  blood  vessels  of  the  organs 
of  the  different  parts  of  the  body  is  also  co-ordinated  by  the  same  regu- 
lative and  controlling  influence  of  the  nervous  system. 

The  Discovery  of  the  Vaso-motor  Nerves. — More  than  half  a  cen- 
tury ago  (1851)  it  was  shown  by  Claude  Bernard  that  if  the  cervical  sym- 
pathetic nerve  is  divided,  the  blood  vessels  of  the  corresponding  side  of  the 
head  and  neck  become  dilated.  This  effect  is  best  observed  in  the  ear, 
which  if  held  up  to  the  light  is  seen  to  become  redder  and  the  arteries  to 
become  larger.  The  whole  ear  is  distinctly  warmer  than  the  opposite  one. 
This  effect  is  produced  by  removing  the  arteries  from  the  tonic  influence  of 
the  central  nervous  system,  which  influence  normally  passes  along  the  course 
of  the  divided  nerve. 

If  the  peripheral  end  of  the  divided  nerve  be  stimulated  in  its  course 
toward  the  organ,  i.e.,  that  farthest  from  the  brain,  the  arteries  which  were 
before  dilated  return  to  their  natural  size,  and  the  parts  regain  their  former 
condition.  And,  besides,  if  the  stimulus  is  very  strong  or  very  long-continued, 
the  amount  of  normal  constriction  is  passed  and  the  vessels  become  much 
more  contracted  than  before.  The  natural  condition,  which  is  midway 
between  extreme  contraction  and  extreme  dilatation,  is  called  the  natural 


244 


THE    CIRCULATION    OF    THE  BLOOD 


tone  of  an  artery.  If  this  is  not  maintained,  the  vessel  is  said  to  have  lost 
tone,  or,  if  it  is  exaggerated,  the  tone  is  said  to  be  too  great.  The  effects 
described  as  having  been  produced  by  section  of  the  cervical  sympathetic 
and  by  subsequent  stimulation  are  not  peculiar  to  that  nerve  and  the  vessels 
to  which  it  is  distributed. 


A  B 

FIG.  201. — Small  Artery  and  Vein  of  the  Frog's  Web.  A,  Under  normal  conditions; 
B,  upon  stimulation  of  the  sciatic  nerve;  Ar,  artery;  V,  vein.  In  this  experiment  the  vein 
also  showed  well-marked  vaso-constriction.  (Greene.) 


It  has  been  found  that  for  every  part  of  the  body  there  exists  a  nerve  the 
division  of  which  produces  the  same  effects,  viz.,  dilatation  of  the  vessels. 
Such  may  be  cited  as  the  case  with  the  sciatic,  the  splanchnic  nerves,  and 
the  nerves  of  the  brachial  plexus;  when  these  are  divided,  dilatation  of  the 
blood  vessels  in  the  parts  supplied  by  them  takes  place.  It  appears, 
therefore,  that  nerves  exist  which  have  a  distinct  control  over  the  vascular 
supply  of  every  part  of  the  body.  These  are  called  vase-motor  or  vaso- 
constrictor nerves.  But  the  arterioles  are  also  under  the  influence  of  a 
second  set  of  nerves,  also  discovered  by  Claude  Bernard,  which  produce 
exactly  the  opposite  influence,  i.e.,  dilatation.  These  nerves  are  called  vaso- 
dilator nerves. 

Mall  has  also  shown  that  veins,  at  least  the  postal  vein,  possess  a  vaso- 
motor  nerve  supply  as  well  as  arteries. 

Vaso-constrictor  Nerves. — The  presence  of  vaso-constrictor  nerves 
can  be  shown  in  several  different  ways,  of  which  the  most  convincing  is  that 
of  direct  inspection.  If  a  vascular  membrane,  like  the  web  of  the  frog's 


VASO-CONSTRICTOR    NERVES 


245 


foot  or  the  bat's  wing,  be  adjusted  on  the  stage  of  a  microscope  for  direct 
inspection,  and  the  smaller  arterioles  are  under  observation,  then  upon  the 
stimulation  of  the  general  nerve  supplying  the  part  these  arterioles  will 
sharply  decrease  in  size.  In  fact,  the  vaso-constriction  is  often  so  great  as  com- 
pletely to  occlude  the  vessel.  Very  soon  after  the  stimulation  the  vessel 
again  dilates  to  its  normal  size. 

The  presence  and  course  of  the  vaso-constrictor  nerve  supply  to  the 
organs  of  the  body  have  been  demonstrated  not  by  direct  inspection,  but 
by  the  use  of  various  forms  of  the  plethysmograph.  A  plethysmograph  is  an 


FIG.  202. — Arm  Plethysmograph.  Apparatus  for  measuring  the  change  in  volume  in 
the  arm  due  to  variation  in  the  blood  supply.  The  arm  is  enclosed  in  a  glass  cylinder 
which  is  completely  filled  with  fluid,  the  opening  through  which  the  arm  is  inserted  being 
closed  by  a  rubber  sleeve,  A.  The  cavity  of  the  glass  cylinder  communicates  through  the 
tube,  F,  G,  with  the  test-tube,  M,  which  is  supported  in  the  jar,  P.  Any  variation  in 
volume  in  the  arm  will  cause  water  to  flow  out  or  into  the  test-tube,  M,  which  is  lowered 
as  the  tube  fills,  and  raised  as  it  empties.  The  rise  and  fall  of  the  test  tube,  M,  is  com- 
municated over  the  pulley,  L,  to  the  writing  pen,  N,  which  records  the  movements  on  the 
smoked  cylinder.  Kymograph  not  shown.  (Mosso.) 

instrument  designed  to  measure  the  variations  in  the  volume  of  an  organ. 
If  the  finger,  the  whole  hand,  the  spleen,  or  the  kidney  be  placed  in  such  an 
instrument  and  the  proper  steps  be  taken  to  record  the  volume  changes,  it 
will  be  found  that  the  volume  of  the  enclosed  organ  is  constantly  changing 
with  every  variation  of  the  blood-pressure.  If  the  nerves  to  the  organ  are 


246  THE    CIRCULATION    OF   THE   BLOOD 

stimulated  by  the  usual  rapidly  interrupted  induction  current,  for  example 
the  splanchnics  to  the  kidney,  then  there  is  a  decrease  in  the  volume  of  the 
organ.  This  decrease  takes  place  even  when  there  is  a  simultaneous  in- 
crease of  the  arterial  blood-pressure,  a  result  that  can  be  explained  only  on 
the  assumption  of  vascular  decrease  in  the  organ.  The  decrease  in  the  flow 
of  blood  to  the  specific  organ  can  be  induced  only  by  a  great  decrease  in  the 
size  of  the  arterioles  produced  by  contractions  of  the  circular  muscles  of 
their  walls. 


a  b 

FIG.  203. — Plethysmograms  of  the  Hind  Leg  of  the  Cat,  showing  a,  vaso-contriction 
on  stimulating  the  sciatic  at  the  rate  of  15  stimuli  per  second  for  twenty  seconds.  In  b, 
the  dilation  of  the  blood  vessels  of  the  opposite  leg  of  the  same  animal  is  shown  on 
stimulating  the  sciatic  which  had  been  cut  four  days  previously.  The  vaso-con- 
strictor nerves  were  degenerated  and  the  vaso-dilators  still  active.  (Bowditch  and 
Warren.) 

Vaso-motor  Tone. — Vaso-constrictor  changes  are  constantly  occur- 
ring in  the  blood  vessels  of  the  organs  of  the  body,  a  fact  that  has  been 
abundantly  demonstrated  by  the  plethysmographic  experiments  just  men- 
tioned. Direct  inspection  of  the  ear  of  an  albino  rabbit  will  show  that  the 
arteries,  and  veins  as  well,  are  now  full  and  large  and  red,  and  the  inter- 
spaces filled  with  blood,  and  now  pale  and  constricted,  and  the  interspaces 
relatively  bloodless.  If  the  cervical  sympathetic  is  cut  as  in  Bernard's 
experiment,  then  the  ear  vessels  remain  dilated,  that  is  they  lose  their 
tone.  This  shows  that  the  condition  is  dependent  primarily  on  the  con- 
stant discharges  of  nerve  impulses  from  the  nervous  system.  It  is  said 
that  the  vessels  regain  their  tone  after  a  time  when  the  nerves  are  cut. 
The  regained  power  may  be  ascribed  to  the  muscle  fibers  themselves. 

Vaso-constrictor  Center. — When  the  tonic  influence  exerted  by  the 
nerve  fibers  on  the  arterioles  is  traced  back  into  the  central  nervous  system, 
it  is  found  to  be  associated  with  the  activity  of  certain  groups  of  nerve  cells, 
or  centers,  which  are  called  the  vaso-constrictor  centers.  This  determi- 
nation is  made  in  part  by  the  method  of  sectioning.  A  lesion  of  the 
cerebro-spinal  axis  below  the  corpora  quadrigemina  is  followed  by  partial 
or  complete  general  dilatation  of  the  blood  vessels  and  a  great  fall  of  blood- 
pressure.  This  is  due  to  the  isolation  of  the  vaso-constrictor  center,  which 
lies  in  the  floor  of  the  fourth  ventricle,  a  few  millimeters  caudal  to  the 
corpora  quadrigemina,  and  extends  longitudinally  over  an  area  of  about 
three  millimeters.  Owsjannikow  has  shown  that  the  center  is  composed  of 


VASO-CONSTRICTOR    REFLEXES 


247 


|Aor, 


VKC 


two  halves,  each  half  lying  in  the  lateral  column  to  the  side  of  the  median 
line.  This  center  is  in  constant  action  during  life,  and  its  discharges  are 
responsible  for  the  vascular  tone  described  in  the  previous  paragraph. 
The  vaso-constrictor  center  varies  in  its  activity,  sometimes  producing 
wave-like  contractions  with  relaxations  of  the  arterial  walls,  producing 
variations  in  the  blood-pressure  known  as  Traube- 
Hering  waves.  These  waves  are  more  of  ten  observed 
in  mammalian  blood-pressure  experiments  after  pro- 
longed operations,  when  the  center  may  be  supposed 
to  be  itself  in  a  weakened  condition. 

Secondary  vaso-motor  centers  are  present  in  the 
spinal  cord  as  proven  by  Goltz.  Under  normal  con- 
ditions they  do  not  act  independently  of  the  medul- 
lary center;  but  when  the  function  of  the  latter  has 
been  interrupted  by  section  of  the  cord,  then  after 
a  few  days  the  spinal  cells  below  the  section  take 
on  central  functions  and  bring  about  a  re-establish- 
ment of  the  lost  vascular  tone.  If  these  centers  be 
destroyed  by  the  destruction  of  the  cord,  then  the 
tone  of  the  vessels  immediately  disappears,  but  is 
regained  after  the  lapse  of  a  much  longer  time.  This 
can  be  ascribed  to  the  presence  of  possible  sympa- 
thetic constrictor  centers  or  more  probably  to  a 
fundamental  property  of  the  muscles  themselves. 
This  experiment  was  carried  out  by  Goltz  and 
Oswald,  who  found  that  after  destruction  of  the 
lower  part  of  the  spinal  cord,  the  tone  of  the  vessels  along  the  Cervical  Sym- 
of  the  hind  limbs,  lost  as  a  result  of  the  operation,  §££££5  SpUnchnic. 
was  later  re-established.  Aur.  Artery  of  ear;  G. 

Vaso-constrictor  Reflexes. — Under  normal  con-  gangiion^^w.  F,  au- 
ditions the  medullary  center  responds  to  afferent  nulus  of  Vieussens;  G. 
stimuli  by  vaso-motor  reflexes.  The  secondary  vaso-  j '  ^jf  pV  thoracic 
motor  centers  in  the  spinal  cord,  when  removed  from  spinal  nerves;  Abd.  Spl, 
the  influence  of  the  bulbar  center,  can  and  do  ^he^rows  indicate  the 
respond  to  afferent  impulses  by  similar  vaso-motor  direction  of  vaso-con- 
strictor impulse, 
action. 

The  afferent  impulses  which  excite  reflex  vaso-motor  action  may  pro- 
ceed from  the  terminations  of  sensory  nerves  in  general,  or  possibly  from 
the  blood  vessels  themselves,  and  the  constriction  which  follows  generally 
occurs  in  the  area  whence  the  impulses  arise.  Yet  the  reflex  may  appear 
elsewhere.  Impulses  proceeding  to  the  vaso-motor  center  from  the  cere- 
brum may  cause  vaso-dilatation,  as  in  blushing,  or  vaso-constriction,  as  in 
the  pallor  of  fear  or  of  anger. 


FIG.      204. — Diagram 


248 


THE    CIRCULATION   OF   THE   BLOOD 


Afferent  influence  upon  the  vaso-motor  centers  is  well  shown  by  the 
action  of  the  depressor  nerve,  the  existence  of  which  was  demonstrated  by 
Cyon  and  Ludwig.  The  depressor  is  a  small  afferent  nerve  which  passes 
up  to  the  medulla  from  the  heart,  in  which  it  takes  its  origin.  It  runs  up- 
ward in  the  sheath  of  the  vagus  or  in  the  superior  laryngeal  branch  of  the 
vagus  or  as  an  independent  branch,  as  in  the  rabbit,  communicating  by 
filaments  with  the  inferior  cervical  ganglion  as  it  proceeds  from  the  heart. 
If,  in  a  rabbit,  this  nerve  be  divided  and  the  central  end  stimulated  during 
a  blood-pressure  observation,  a  remarkable  fall  of  blood-pressure  takes 
place,  figure  205. 


FIG.  205. — Blood-pressure  Record  (lower)  and  Respiratory  Record  (upper)  Obtained 
from  a  Dog  upon  Stimulating  the  Central  End  of  the  Divided  Vagus,  Both  Vagi  being  Cut. 
The  marked  fall  in  blood -pressure  is  due  to  the  effect  of  stimulating  the  depressor  fibers 
contained  in  the  vagus  trunk  of  the  dog.  (New  figure  by  Dooley  and  Dandy.) 

The  cause  of  the  fall  of  blood  pressure  is  found  to  proceed  primarily 
from  the  dilatation  of  the  vascular  district  within  the  abdomen  supplied  by 
the  splanchnic  nerves,  in  consequence  of  which  the  vessels  hold  a  much 
larger  quantity  of  blood  than  usual.  The  engorgement  of  the  splanchnic 
area  very  greatly  diminishes  the  amount  of  blood  in  the  vessels  elsewhere, 
and  so  materially  diminishes  the  blood-pressure.  The  function  of  the  de- 
pressor nerve  is  that  of  conveying  to  the  vaso-motor  center  afferent  nerve 
impulses  from  the  heart,  which  produce  an  inhibition  of  the  tonic  activity 
of  the  vaso-motor  center  and,  therefore,  a  diminution  of  the  tension  in  the 
blood  vessels.  This  diminishes  the  overstrain  on  the  heart  in  propelling 
blood  into  the  already  too  full  or  too  tense  arteries.  It  has  been  shown  by 
Porter  and  Beyer  that  the  fall  in  blood-pressure,  following  stimulation  of  the 
depressor  nerve,  will  still  occur,  even  when  the  abdominal  vaso- constriction 
is  kept  constant  by  a  simultaneous  stimulation  of  the  splanchnics.  It  is 
therefore  evident  that  the  inhibitory  effect  of  depressor- nerve  stimulation  is 
a  general  one  and  not  confined  to  the  splanchnic  area  alone. 


VASO-DILATOR    NERVES  249 

The  action  of  the  depressor  nerve  in  causing  an  inhibition  of  the  vaso- 
motor  center  illustrates  the  more  unusual  effect  of  afferent  impulses;  that  is, 
inhibition  of  the  vaso-constrictor  tone.  As  a  rule,  the  stimulation  of  the 
central  end  of  an  afferent  nerve,  such  as  the  sciatic  or  the  internal  saphenous, 
produces  the  reverse,  i.e.,  a  pressor  effect,  and  increases  the  tonic  influence 
of  the  center  which  by  causing  constriction  of  the  arterioles  raises  the  blood 
pressure.  Thus  the  reflex  effects  of  stimulating  an  afferent  nerve  may  be 
either  to  constrict  or  to  dilate  the  arteries.  These  reflexes  may  be  general 
enough  to  influence  the  general  blood-pressure  or  they  may  be  limited  to 
definite  local  areas,  but  the  local  effects  are  the  all-important  ones,  since 
by  these  the  local  regulation  of  the  blood  flow  is  accomplished. 

Traube-Hering  Curves. — The  vaso-motor  center  sends  out  rhythmical 
impulses  by  which  undulations  of  blood-pressure  of  a  large  and  sweeping 
character  are  produced,  quite  independent  of  the  so-called  respiratory  un- 
dulations. The  action  of  this  center  in  producing  such  undulations  is  dem- 
onstrated in  the  following  observations.  In  an  animal  under  the  influence 
of  curare  and  with  both  vagi  cut,  and  a  record  of  whose  blood-pressure  is 
being  taken,  if  artificial  respiration  be  stopped,  the  blood-pressure  rises 
sharply  at  first.  After  a  time  the  rhythmical  undulations  shown  in  figure  206 
occur.  These  variations  are  called  Traube's  or  Traube-Hering  curves. 
There  may  be  upward  of  ten  of  the  respiratory  undulations  in  one  Traube- 
Hering  curve.  They  continue  until  the  vaso-motor  center  is  asphyxiated 
and  the  heart  exhausted,  when  the  pressure  falls.  The  undulations  cannot 
be  caused  by  anything  but  the  vaso-motor  center,  as  the  mechanical  effects 
of  respiration  have  been  eliminated  by  the  curare  and  by  the  cessation  of 
artificial  respiration,  and  the  effect  of  the  cardio-inhibitory  center  has  been 
removed  by  the  division  of  the  vagi.  The  rhythmic  rise  of  blood-pressure 
is  most  likely  due  to  a  rhythmic  constriction  of  the  arterioles  followed  by  a 
corresponding  relaxation  and  fall  of  pressure,  both  being  due  to  the  action 
of  the  vaso-motor  center.  The  vaso-motor  center,  therefore,  is  capable  of 
producing  rhythmical  discharges  of  vaso-constrictor  nerve  impulses  that 
result  in  the  undulations  of  blood-pressure. 

Vaso-dilator  Nerves. — Claude  Bernard  discovered  (1856)  that  the 
blood  flow  was  increased  through  the  salivary  glands  by  stimulation  of  the 
nerves  (the  chorda  tympani  for  the  submaxillary,  and  the  tympanic  branch 
of  the  glossopharyngeal  for  the  parotid),  thus  proving  that  the  arteries  have 
not  only  vaso-constrictors,  but  also  vaso-dilator  nerves.  Vaso-dilator  nerves 
have  been  described  for  most  parts  of  the  body.  In  general  they  are  dis- 
tributed in  the  same  nerve  trunks  which  bear  the  vaso-constrictors. 

It  is  not  supposed  that  the  vaso-dilators  produce  widening  of  the 
arterioles  by  stimulation  to  active  muscular  contraction;  in  fact,  the  circular 
arrangement  of  the  muscle  fibers  would  seem  to  exclude  such  a  deduction. 
It  is  probable  that  there  is  local  inhibition  of  the  tonic  contraction  of  the 


250  THE  CIRCULATION  OF  THE  BLOOD 

muscles,  thus  allowing  the  mechanical  factor  of  the  general  blood-pressure 
to  dilate  the  vessels.  The  vaso-dilator  nerves  are  characterized  by  their  re- 
sponse to  slowly  developed  stimuli,  shown  by  Bowditch  and  Warren, 
and  by  the  retention  of  irritability  after  degeneration  of  the  constrictors 
has  taken  place,  see  figure  203. 


FIG.  206. — Traube-Hering's  Curves.  (To  be  read  .from  left  to  right.)  The  curves 
i,  2,  3,  4,  and  5  are  portions  selected  from  one  continuous  tracing  forming  the  record  of  a 
prolonged  observation,  so  that  the  several  curves  represent  successive  stages  of  the  same 
experiment.  Each  curve  is  placed  in  its  proper  position  relative  to  the  base  line,  which  is 
omitted;  the  blood-pressure  rises  in  stages  from  i  to  2,  3,  and  4,  but  falls  again  in  stage  5. 
Curve  i  is  taken  from  a  period  when  artificial  respiration  was  being  kept  up,  but,  the  vagi 
having  been  divided,  the  pulsations  on  the  ascent  and  descent  of  the  undulations  do  not 
differ;  when  artificial  respiration  ceased,  these  undulations  for  a  while  disappeared,  and  the 
blood-pressure  rose  steadily  while  the  heart-beats  became  slower.  Soon,  as  at  2,  new 
undulations  appeared;  a  little  later,  the  blood-pressure  was  still  rising,  the  heart- beats  still 
slower,  but  the  undulations  still  more  obvious  3;  still  later  4,  the  pressure  was  still 
higher,  but  the  heart-beats  were  quicker,  and  the  undulations  flatter;  the  pressure  then 
began  to  fall  rapidly  5,  and  continued  to  fall  until  some  time  after  artificial  respiration  was 
resumed.  (M.  Foster.) 


Vaso-dilator  Centers.— No  distinct  medullary  center  has  yet  been  shown 
to  regulate  the  vaso-dilator  nerve  activity.  Such  centers,  if  they  exist, 
should  be  influenced  by  isolating  them  from  their  efferent  paths,  on  the  one 
hand,  or  by  stimulation  by  afferent  channels,  on  the  other.  The  former 
method  of  study  has  revealed  nothing  that  can  be  compared  to  the  tonic  ac- 


VASO-DILATOR    REFLEXES 


251 


tivity  of  the  constrictor  center.  Efferent  dilator-nerve  impulses  can  be  re- 
flexly  produced  by  sensory  stimulation.  The  isolated  lumbar  cord  of  a  dog 
is  capable  of  reflex  vaso-dilator  activity,  since  stimulation  of  the  skin  of 
the  penis  leads  to  reflex  vaso-dilatation,  indicating  the  presence  of  local 
vaso-dilator  reflex  mechanisms  or  paths  through  the  lumbo-sacral  portion 
of  the  spinal  cord. 

TABLE  SHOWING  THE  INFLUENCE  OF  RATE  AND  STRENGTH  OF  STIMULATION  OF  THE 
SCIATIC  NERVE  ON  THE  BLOOD  VESSELS  OF  THE  LEG  (BOWDITCH  AND  WARREN) 


Weak  Ind.  «»  10-19 

Medium  Ind.   =    100-250 

Strong  Ind.    =    300-800 

Stimu- 

rate  per 
second 

No.  of 
obs. 

Contr. 
mm. 

Dilat. 

mm. 

No.  of 
obs. 

Contr. 
mm. 

Dilat. 
mm. 

No.  of 
obs. 

Contr. 

mm. 

Dilat. 

mm. 

I/IO-I 

45 

I  .  2 

6.2 

146 

i  .9 

4-5 

85 

2.6 

32 

2-4 

63 

3-6 

5-6 

72 

4-4 

5-4 

57 

7.6 

6.4 

5-8 

20 

4-5 

4-5 

49 

6-3 

3-2 

75 

8.7 

40 

10-16 

49 

10.3 

2.4 

104 

7-8 

2.8 

34 

n.  4 

2.7 

30-64 

62 

14-3 

2.  I 

40 

17.6 

3-1 

8 

14.6             1.4 

1 

Vaso-dilator  Reflexes. — Perhaps  the  only  unquestioned  case  of  reflex 
vaso-dilatation  is  that  of  the  lumbar  cord  just  mentioned.  It  is  true  that 
many  apparent  reflexes  can  be  noted,  for  example  the  increased  flow  of 
blood  in  the  salivary  glands  under  gustatory  reflexes,  the  blushing  of  the 
skin  on  exposure  to  sudden  warmth,  or  even  the  blushing  of  emotional 
origin.  These  on  first  thought  would  be  regarded  as  vaso-dilator  reflexes. 
In  all  these  cases  there  is  a  widening  of  the  peripheral  arterioles  with  a 
great  increase  in  the  volume  of  blood  flowing  through  the  local  vascular 
bed.  But  each  instance  can  be  just  as  readily  explained  as  inhibition 
of  the  vaso-constrictor  tonic  activity.  This  double  explanation  can,  as  a 
matter  of  fact,  be  applied  to  the  action  of  the  depressor  nerve  described 
above,  page  248.  However,  the  confusion  is  in  part  due  to  the  diffi- 
culty of  analyzing  the  two  classes  of  nerves  in  the  same  nerve  trunk. 
All  the  thoracic  spinal  nerves  and  the  upper  lumbar  trunks  contain  both 
vaso-constrictor  and  vaso-dilator  nerves.  The  usual  methods  of  physio- 
logical stimulation  with  rapidly  interrupted  currents  we  now  know  are 
normal  stimuli  for  the  constrictors  only.  Stimulations  at  the  rate  of  one 
or  two  per  second  call  forth  responses  in  the  dilator  fibers.  Also,  the  vaso- 
dilator fibers  degenerate  more  slowly  and  retain  their  irritability  longer 
than  the  constrictor  fibers  when  both  are  isolated  from  their  cell  bodies 
by  sectioning.  The  method  of  slow  stimulation  and  the  method  of  differ- 
ential degeneration  were  given  us  by  Bowditch  and  Warren. 


252  THE    CIRCULATION   OF   THE   BLOOD 

The  Relation  of  Vaso-constrictor  and  Vaso-dilator  Activity. — The 
distribution  of  two  sets  of  regulative  fibers  for  the  muscular  walls  of  the 
blood  vessels,  when  considered  in  connection  with  the  other  factors  of  the 
vascular  apparatus,  gives  a  wonderfully  complete  mechanism  for  the  co- 
ordination of  the  vascular  supply  with  the  activity  of  the  different  organs. 
General  and  broadly  distributed  activity  of  the  constrictors  produces 
increase  of  general  blood  pressure  and  of  the  dilators  decrease  of  pressure, 
but  local  activity  of  either  set  will  produce  a  great  reduction  or  increase  of 
blood  in  the  local  organ  with  little  or  no  effect  on  the  general  pressure. 
When  a  vaso-dilatation  is  produced  locally  in  one  organ  and  there  is  an 
accompanying  vaso-constriction  in  other  regions,  as  usually  happens,  it  is 
evident  that  the  result  may  be  a  flooding  of  the  local  region.  This  is 
exactly  the  thing  that  is  accomplished  in  the  muscles  in  violent  exercise, 
in  the  glands  during  secretion,  in  the  stomach  during  digestion.  It  is 
this  mechanism  that  is  utilized  to  throw  a  large  volume  of  blood  to  the 
skin  when  the  temperature  of  the  body  is  above  the  average,  or  to  blanch 
the  skin  when  the  temperature  is  low. 

Normally  certain  regions  of  the  body  are  associated  in  that  when  vaso- 
dilatation  occurs  n  one  region,  vaso-constriction  occurs  in  the  other.  This  is 
particularly  true  with  the  skin  or  surface  of  the  body  and  the  viscera  or  deeper 
organs.  The  same  relation  is  said  to  exist  between  some  of  the  visceral 
organs. 

General  Course  of  the  Vaso-constrictor  and  Vaso-dilator  Nerves.— 
The  cell  bodies  forming  the  medullary  vaso-motor  center  give  off  axones, 
axis- cylinder  processes,  some  of  which  go  to  the  nuclei  of  origin  of  certain 
cranial  nerves,  while  others  pass  down  the  cord  to  end  at  different  levels 
in  contact  with  certain  cells,  probably  small  cells  in  the  anterior  horn  and 
lateral  part  of  the  gray  matter.  These  cord  cells  constitute  the  spinal  centers. 
The  neuraxones  of  the  spinal  cells  leave  the  cord  in  certain  spinal  nerves  in 
the  anterior  roots,  pass  by  the  white  rami  to  the  sympathetic  ganglionic 
chain,  where  they  end  in  physiological  connection  with  the  ganglionic  cell. 
Axones  from  these  latter  cells  pass  by  an  uninterrupted  course  to  their  termina- 
tions on  the  blood  vessel  walls.  The  vaso-constrictor  fibers  leave  the  central 
nervous  axis  by  the  ventral  roots  of  all  the  dorsal  nerves  and  the  first  two 
lumbar  roots,  a  comparatively  restricted  region.  The  vaso-dilators  have 
the  same  origin  with  two  exceptions,  viz.,  the  vaso-dilators  of  the  salivary 
glands  found  in  the  seventh  and  ninth  cranial  nerves,  and  the  nervi  erigentes, 
which  arise  in  the  roots  of  the  second  and  third  sacrals. 

The  nerves  to  the  viscera  pass  direct  to  their  blood  vessels,  but  the  vas- 
cular nerves  for  the  skin,  muscles,  limbs,  etc.,  rejoin  the  main  divisions  of  the 
spinal  nerves  through  the  gray  rami,  see  figures  417  and  418,  and  pass  to  the 
blood  vessels  along  with  the  general  nerves  of  the  organ  or  organs. 


VASO-CONSTRICTOR    AND   VASODILATOR    NERVES  253 

VASO-CONSTRICTOR  AND  VASO-DILATOR  NERVES  FOR 
INDIVIDUAL  ORGANS. 

The  particular  paths  for  the  vaso-motor  nerves  has  been  pretty  definitely 
established  by  numerous  researches,  especially  by  those  of  Langley  and  his 
students. 

The  course  of  the  vaso-constrictor  and  the  vaso-dilator  nerve  fibers  has 
been  followed  satisfactorily  in  many  of  the  important  parts  of  the  body, 
though  the  supply  for  some  regions  is  yet  obscure.  This  is  particularly 
true  for  the  brain,  where  such  supply  is  apparently  absent.  The  two  groups 
of  fibers  run  the  same  course,  except  in  the  cephalic  and  sacral  regions  already 
mentioned  They  may,  therefore,  be  described  together. 

The  Vascular  Nerve  Supply  for  the  Head.— The  vascular  nerves 
for  the  head,  face,  and  mouth  have  their  origin  in  the  cord  from  the  first  to 
the  fifth  dorsal  spinal  nerves.  They  pass  through  the  white  rami  to  sym- 
pathetic ganglia,  through  the  stellate  ganglion,  and  up  the  cervical  sympa- 
thetic nerve  to  the  superior  cervical  ganglion.  From  this  ganglion  they  run 
to  their  distribution,  either  along  with  the  arteries,  as  with  the  salivary  sup- 
ply, or  with  the  sensory  nerves,  as  in  the  nerves  to  the  mucous  membrane 
of  the  mouth,  etc.  The  vascular  nerves  supplied  to  the  base  of  the  ear  follow 
the  above  course,  but  the  nerves  for  the  tip  leave  the  stellate  ganglion  in  the 
ramus  vertebralis,  run  to  the  third  cervical  nerve,  and  pass  with  its  auricular 
branch  to  the  ear,  a  circuitous  route  determined  by  Fletcher. 

The  great  exception  to  the  above  origin  is  with  the  vaso-dilator  group. 
Dilator  fibers  leave  the  base  of  the  brain  in  the  direct  path  of  the  seventh 
cranial  nerve  to  supply  the  submaxillary  and  sublingual  glands,  in  the  ninth 
cranial  nerve  to  the  parotid  gland,  and  in  both  these  to  the  tongue. 

The  Vascular  Regulation  in  the  Brain. — The  brain  requires  a  large 
and  uniform  supply  of  blood  for  the  due  performance  of  its  functions.  This 
object  is  effected  through  the  number  and  size  of  its  arteries,  the  two  internal 
carotids  and  the  two  vertebrals.  It  is  also  desirable  that  the  force  with 
which  this  blood  is  sent  to  the  brain  should  be  subject  to  less  variation  from 
external  circumstances  than  it  is  in  other  parts,  an  effect  that  is  accomplished 
by  the  free  anastomoses  of  the  large  arteries  in  the  circle  of  Willis.  This 
arrangement  insures  that  the  supply  of  blood  will  be  uniform  in  both  hemi- 
spheres even  though  it  may^be  limited  through  operation  or  accident  to  one 
or  more  of  the  four  principal  arteries.  Uniformity  of  supply  is  further 
insured  by  the  arrangement  of  the  vessels  in  the  pia  mater.  Previous  to 
their  distribution  to  the  substance  of  the  brain  the  large  arteries  break  up 
and  divide  into  innumerable  minute  branches.  These  arterioles  after 
frequent  communication  with  one  another  enter  the  brain  in  a  very  uniform 
and  equable  distribution.  The  arrangement  of  the  veins  within  the  cranium 
is  also  peculiar.  The  large  venous  trunks  or  sinuses  are  formed  so  as  to  be 
scarcely  capable  of  change  of  size;  and  composed,  as  they  are,  of  the  tough 
tissue  of  the  dura  mater,  and  in  someinstancesbounded  by  the  bony  cranium, 


254  THE    CIRCULATION    OF   THE   BLOOD 

they  are  not  compressible  by  any  force  which  the  fullness  of  the  arteries 
might  exercise  through  the  substance  of  the  brain.  Nor  do  they  admit  of 
distention  when  the  flow  of  venous  blood  from  the  brain  is  obstructed. 

The  mechanical  conditions  in  the  brain  and  skull  formerly  appeared 
enough  to  justify  the  opinion  that  the  quantity  of  blood  in  the  brain  must 


FIG.  208. — Showing  the  Origin  and  Course  of  the  Vaso -constrictor  Nerves  for  the 
Head.  M,  medulla;  C8,  eighth  cervical  spinal  nerve;  V,  vagus,  S.c.g.,  superior  cervical 
ganglion.  Modified  from  Moret. 

be  at  all  times  the  same.  But  it  was  found  that  in  animals  bled  to  death 
without  any  aperture  being  made  in  the  cranium,  the  brain  became  pale  and 
anemic  like  other  parts.  And  in  death  from  strangling  or  drowning,  there 
was  congestion  of  the  cerebral  vessels;  while  in  death  by  prussic  acid,  the 
quantity  of  blood  in  the  cavity  of  the  cranium  was  determined  by  the  position 
in  which  the  animal  was  placed  after  death,  the  cerebral  vessels  being  con- 
gested when  the  animal  was  suspended  with  its  head  downward,  and  com- 
paratively empty  when  the  animal  was  kept  suspended  by  the  ears.  Thus, 
although  the  total  volume  of  the  contents  of  the  cranium  is  probably  nearly 
always  the  same,  yet  the  quantity  of  blood  in  it  is  liable  to  variation,  its  in- 
crease or  diminution  being  accompained  by  a  simultaneous  diminution  or 
increase  in  the  quantity  of  the  cerebro-spinal  fluid.  The  cerebro-spinal 
fluid  being  readily  removed  from  one  part  of  the  brain  and  spinal  cord  to 


VASCULAR    REGULATION    IN    THE   BRAIN  255 

another,  and  capable  of  being  rapidly  absorbed  and  as  readily  effused,  would 
serve  as  a  kind  of  supplemental  fluid  to  the  other  contents  of  the  cranium 
to  keep  it  uniformly  filled.  Although  the  arrangement  of  the  blood  vessels 
insures  to  the  brain  an  amount  of  blood  which  is  tolerably  uniform,  yet  with 
every  beat  of  the  heart,  and  every  act  of  respiration,  and  under  many  other 
circumstances,  the  quantity  of  blood  in  the  cavity  of  the  cranium  is  con- 
stantly varying.  Roy  and  Sherrington  are  responsible  for  the  view  now 
generally  current  that  the  brain,  therefore,  is  largely  if  not  entirely  dependent 
upon  the  general  blood-pressure  for  variations  in  the  quantity  of  blood  which 
it  receives.  During  a  high  blood-pressure  the  amount  of  blood  that  flows 
in  a  given  unit  of  time  is  greater,  and  during  low  blood-pressure  less.  Howell 
has  shown  that  in  the  decapitated  dog's  brain  the  flow  of  blood  is  directly 
proportional  to  the  difference  in  pressure. 

Numerous  attempts  have  been  made  to  show  vaso-motor  mechanisms 
for  the  cerebral  arteries,  but  with  generally  unconvincing  success.  Huber 
and  others  have  shown  nerve  endings  in  such  arteries  by  histological 
methods.  Bayless,  Hill,  and  Gulland  make  the  statement  that  "no- 
evidence  has  been  found  of  the  existence  of  cerebral  vaso-motor  nerves, 
either  by  means  of  stimulation  of  the  vaso-motor  center  or  central  end  of 
the  spinal  cord,  after  division  of  the  cord  in  the  upper  dorsal  region,  or 


FIG.  209. — Vaso-dilatation  in  the  Brain  from  Stimulation  of  the  Cerebral  Cortex 
in  the  Presence  of  Complete  Destruction  of  the  Medulla  in  the  Cat.  The  upper  trace 
is  of  the  carotid  pressure;  the  lower  trace  is  of  the  brain  oncometer.  (Weber.) 

by  stimulation  of  the  stellate  ganglion,  and  that  is  to  say  the  whole 
sympathetic  supply  to  the  carotid  and  vertebral  arteries."  However, 
Ernest  Weber  (1908)  reinvestigated  the  control  of  the  blood  flow  in  the 
brain.  He  admits  that  the  blood  flow  in  the  brain  is  sharply  dependent 
on  the  general  blood-pressure,  but  he  presents  plausible  evidence  that 
both  vaso-constrictors  and  vaso-dilators  exist  for  the  brain  vessels.  The 
most  striking  facts  are  obtained  upon  stimulating  general  sensory  nerves, 
the  central  end  of  the  sectioned  cord,  the  cerebral  cortex,  and  the  cervical 
sympathetic.  The  stimulation  of  the  cerebral  cortex  calls  forth  vaso- 
dilatation  in  the  brain  even  when  the  medulla  is  completely  destroyed. 
Weber,  therefore,  concludes  that  these  stimuli  act  reflexly  through  a 
cerebral  vascular  center  located  at  some  as  yet  undetermined  point  in 
the  brain  stem  above  the  general  medullary  center.  An  active  cerebral 
vaso-dilatation  may  be  accomplished  through  this  center  even  in  the 


THE    CIRCULATION    OF   THE   BLOOD 


presence  of  an  accompanying  fall  in  general  blood-pressure.  More  often 
the  type  of  vascular  reflex  is  that  of  dilatation  followed  by  constriction 
of  the  brain  vessels.  If  there  is  an  accompanying  sharp  rise  in  general 
blood-pressure,  then  the  reflex  cerebral  vascular  dilatation  is  followed  by 


nee.    ur    VES. 


3-a 
| -a. 

u   a) 

S^ 


irl'&iig'^  i 


vaso-constriction  which  takes  place  while  the  general  blood-pressure  is  yet 
high.  Weber  suggests  that  this  local  vaso-constrictor  mechanism  provides 
an  apparatus  for  the  regulation  and  control  of  the  cerebral  congestion 
that  results  from  the  rise  of  general  blood-pressure  due  to  the  operation  of 
the  general  medullary  vaso-motor  center. 


THE   VASCULAR    NERVES  257 

The  question  may  be  summarized  by  the  statement  that  the  regulation 
of  the  flow  of  blood  through  the  brain  is  accomplished  by  the  interaction  of 
two  factors:  First,  the  indirect  regulation  of  blood-pressure  through  the 
variations  in  the  heart's  activity,  and  through  the  general  action  of  the 
medullary  vaso- motor  apparatus  producing  vaso-constrictions  or  dilatations 
in  areas  other  than  the  brain.  Second,  the  local  and  direct  regulation  of 
the  brain  vessels  through  reflex  action  on  a  special  local  vascular  center. 

The  Vascular  Nerves  for  the  Thoracic  Viscera.— Numerous  efforts 
have  been  made  to  determine  the  vaso- motor  nerve  supply  for  the  thoracic 
organs,  the  heart  and  lungs.  In  the  heart  the  observation  is  rendered  com- 
plex by  the  fact  of  the  rhythmic  contractions  which  produce  mechanical 
pressure  on  the  coronary  arteries.  Martin,  by  direct  observation  through  a 
lens,  and  Porter,  by  measuring  the  outflow  of  the  coronaries  upon  vagus 
stimulation,  came  to  exactly  opposite  views:  the  former  that  the  vagus  con- 
tained vaso-dilators,  the  latter  that  it  contained  vaso-constrictors.  Still 
other  experiments  have  been  made  to  prove  either  constrictor  or  dilator 
nerves  for  the  coronary  arteries,  but  the  fact  is  still  undetermined. 

The  lesser  circulation  through  the  lungs  has  also  proven  a  difficult  situa- 
tion to  interpret  as  regards  any  nervous  regulation  of  the  pulmonary  arteri- 
oles.  The  evidence,  while  not  conclusive,  is  that  the  vaso-constrictor  supply 
to  the  lungs  is  from  the  third  to  the  fifth  thoracic  nerves,  but  that  the  vaso- 
constriction  produced  is  slight  in  comparison  with  regions  of  the  systemic 
•circulation. 

The  Vascular  Nerves  for  the  Abdominal  Viscera. — The  vaso-con- 
strictors and  the  vaso-dilators  for  the  organs  of  the  abdominal  cavity  have 
a  broad  origin  in  the  cord,  from  the  first  dorsal  to  the  fourth  lumbar  in 
the  dog  and  cat.  The  nerves  pass  to  the  organs  by  the  splanchnic  nerves, 
and  by  the  solar,  celiac,  and  mesenteric  ganglia.  The  vascular  nerves  for 
the  different  organs  may  be  given  in  tabulated  form: 

Vascular  Nerves  for  the  Abdominal  Viscera. 

Organ.                   Spinal  origin  of  the  vascu-  Course  to  the  organ. 

lar  nerves. 

Stomach  and  in-    \     5,  6,  7,  8,  9,  10,  n,  12,  13  D,  /  Splanchnic  nerves  and 

testine.                   (     i  L                                                |  solar  and  celiac  ganglia. 

o   iee                          I    3,  4,  5,  6,  7,  8,  9,  10,  n,  12,  J  Splanchnic     nerves     and 

/       13  D,  i  L                                 |  solar  and  celiac  ganglia. 

T  •  -r^       f    Splanchnic     nerves     and 

Liver 3,  4,  St  6.  7.  8,  9,  10,  nD.  .  < 

solar  and  celiac  ganglia. 


Kidnev  4>  5>  6'  7'  8'  9>  I0'  "'  I2'  I3        Splanchnic    and    celiac 

/       D,  i,  2,  3,  4  L.  \      ganglia. 

Inferior    splanchnic    and 

Pelvic  viscera i,  2,  3,  4!, -J       inferior       mesenteric 

ganglia. 


258  THE  CIRCULATION  OF  THE  BLOOD 

The  Vascular  Nerves  for  the  External  Genital  Organs. — The  vaso- 
dilators for  these  organs  arise  from  the  second  and  third  sacral  nerves  and 
pass  to  the  organs  by  the  nervi  erigentes  and  the  pelvic  plexus.  They  form 
the  second  great  exception  to  the  region  of  general  outflow  of  vascular  nerves. 
The  constrictors,  on  the  other  hand,  arise  in  the  spinal  nerves  from  the  last 
dorsal  and  first  four  lumbar.  They  run  the  same  course  as  given  in  the  table 
for  the  pelvic  viscera. 

The  greatest  variations  in  the  quantity  of  blood  contained  at  different 
times  in  the  external  genital  organs  are  found  in  certain  structures  which 
contain  what  is  known  as  erectile  tissue.  These  organs,  under  ordinary  cir- 
cumstances, are  soft  and  flaccid,  but  at  certain  times  they  receive  an  un- 
usually large  quantity  of  blood,  become  distended  and  swollen  by  it,  and 
pass  into  the  state  termed  erection.  Such  structures  are  the  corpora  cavernosa 
and  corpus  spongiosum  of  the  penis  of  the  male,  and  the  clitoris  in  the  female. 
The  nipple  of  the  mammary  gland  in  both  sexes,  and,  according  to  some 
authors,  certain  nasal  membranes  contain  erectile  tissue. 

The  corpus  cavernosum  of  the  penis,  which  is  the  best  example  of  an 
erectile  structure,  has  an  external  fibrous  membrane  or  sheath.  From  the 
inner  surface  of  the  sheath  numerous  fine  lamellae  project  into  the  cavity, 
dividing  it  into  small  compartments,  like  cells  when  they  are  inflated.  Within 
these  cells  there  is  a  plexus  of  veins  upon  which  the  erectile  property  of  the 
organ  mainly  depends.  The  plexus  consists  of  short  veins  with  very  close 
interlacings  and  anastomoses  with  very  elastic  walls  admitting  of  great  varia- 
tions in  size.  They  collapse  in  the  passive  state  of  the  organ,  but  are  capable 
of  an  amount  of  dilatation  which  exceeds  beyond  comparison  that  of  the 
arteries  and  veins  which  convey  the  blood  to  and  from  them.  The  strong 
fibrous  tissue  lying  in  the  intervals  of  the  venous  plexuses,  and  the  external 
fibrous  membrane  or  sheath  with  which  it  is  connected,  limit  the  distention 
of  the  vessels  and  give  to  the  organ  its  condition  of  tension  and  firmness. 
The  same  general  condition  of  vessels  exists  in  the  corpus  spongiosum 
urethrae,  but  the  fibrous  tissue  around  the  urethra  is  much  weaker  than 
around  the  body  of  the  penis,  while  around  the  glans  there  is  none.  The 
venous  blood  is  returned  from  the  plexuses  by  comparatively  small  veins; 
all  of  which  are  liable  to  the  pressure  of  muscles  where  they  leave  the  penis. 
The  muscles  chiefly  concerned  in  this  action  are  the  erector  penis  and 
accelerator  urinae.  Erection  results  from  the  distention  of  the  venous 
plexuses  by  a  sudden  influx  of  blood  resulting  from  the  action  of  the  nervous 
vascular  reflexes.  It  is  facilitated  by  the  special  muscular  mechanism 
which  prevents  the  outflow  of  blood. 

The  Vascular  Nerves  for  the  Trunk  and  Limbs. — The  skin  and 
muscles  of  the  trunk  receive  their  cutaneous  and  motor  nerves  by  a  seg- 
mental  arrangement  in  which  the  innervation  is  by  bands  corresponding 
with  the  segments  of  the  cord  and  the  spinal  nerves.  It  is  much  the  same 


THE   VASO-CONSTRICTOR    NERVES    FOR    THE   VEINS 


259 


with  the  vascular  nerves;  they  are  distributed  to  the  skin  and  walls  of  the 
trunk  in  the  same  segment  in  which  they  arise.  Langley  says  that  the  suc- 
cessive bands  overlap  somewhat. 

In  the  fore  legs  or  arms  the  vascular  nerves  arise  from  the  first  to  the 
sixth  dorsal  spinal  nerves,  run  to  the  stellate  ganglia,  then  by  the  gray  rami 
back  through  the  ramus  vertebralis  to  join  those  cervical  nerves  that  enter 
into  the  brachial  plexus,  figure  211. 


FIG.  211. — Plan  of  Distribution  of  Vaso-constrictor  Nerves  for  the  Fore  Limbs. 
(Modified  from  Moret.) 

The  nerves  for  the  blood  vessels  of  the  lower  limbs  arise  from  the  tenth 
dorsal  to  the  second  lumbar  nerves.  These  pass  to  the  ganglionic  chain,  and 
gray  rami  are  given  off  which  join  the  lumbo-sacral  plexus  and  run  with  the 
divisions  of  that  nerve  complex  to  their  distribution  in  the  skin  and  muscles. 
Vaso-constrictors  and  vaso-dilators  have  a  common  course  to  the  lower  limbs. 

The  Vaso-constrictor  Nerves  for  the  Veins. — Mall  has  proven  that 
vaso- constrictors  are  present  for  the  portal  vein.  These  fibers  are  present 
in  the  splanchnic  nerves.  Other  evidences  have  been  observed  which 
render  the  view  probable  that  vaso-motors  for  the  veins  in  general  exist. 
Hough,  for  example,  in  an  extended  study  of  the  capillary  pressure  found 
many  variations  which  were  readily  explained  only  on  the  assumption  of 
veno- motor  activity,  see  figure  201. 


260  THE  CIRCULATION  OF  THE  BLOOD 


LABORATORY  EXPERIMENTS  ON  THE  CIRCULATION 

1.  The  Rate  of  the  Human  Heart-beat. — Determine  the  rate  of  the 
heart-beat  per  minute  by  counting  the  radial  pulse,  using  a  watch  for  the 
time.     Make  the  determination  after  sitting  quietly  for  five  minutes. 
Take  the  average  of  at  least  ten  determinations  for  your  own  case.     Deter- 
mine the  heart-rate  under  the  same  conditions  for  as  many  different  per- 
sons as  you  can.     Tabulate  these  rates  to  show  age,  sex,  weight,  and 
height  of  the  different  classes  of  individuals,  and  compute  general  averages 
for  your  sets.     Count  the  rate  in  children  and  in  aged. 

Note  the  effect  on  the  averages  obtained  above  after  lying  down  for 
five  minutes,  after  standing  quietly  for  the  same  time,  and  after  five 
minutes'  brisk  walk.  Tabulate  as  directed. 

Count  the  heart-rate  by  successive  15  second  periods  immediately 
upon  standing  from  the  reclining  position  and  until  the  quarter  minute 
rates  are  constant;  repeat  immediately  after  two  minutes  of  fast  running. 
Tabulate  these  results  and  compare  the  graphs  obtained  from  several 
different  individuals.  This  method  measures  the  character  of  the  vascular 
control  in  certain  clinical  states. 

Count  your  own  heart-rate  at  one-hour  intervals  during  one  entire 
day,  giving  special  attention  to  the  rate  just  before  and  just  after  meals, 
but  in  every  case  make  the  count  on  the  fifth  minute  while  sitting  quietly. 
A  marked  diurnal  variation  will  usually  appear.  Determine  these  rates 
on  several  individuals,  and  tabulate  as  before. 

2.  Human    Cardiogram. — Apply    a    Burdon-Sanderson    cardiograph 
to  the  thorax  over  the  point  between  the  fifth  and  sixth  ribs  of  the  left 
side,  where  the  cardiac  impulse  is  felt  most  distinctly.     Connect  the 
cardiograph  with  a  recording  tambour,  Marey's  form,  adjust  the  tension 
of  the  cardiograph  and  the  pressure  of  the  air  within  the  system,  and  take 
a  tracing  of  the  movements  of  the  lever  of  the  recording  tambour.     The 
recording  cylinder  should  travel  at  the  rate  of  about  two  centimeters  per 
second.     Take  the  time  of  the  movements  of  the  kymograph  by  an  elec- 
tric seconds  magnet.     The  proper  description  should  be  written  on  the 
smoked  paper,  the  paper  removed  carefully  and  the  whole  record  fixed  in 
shellac. 

Count  the  rate  of  the  heart-beat  from  the  record.  Compute  the  time 
of  the  cardiac  systole  and  diastole,  and  of  the  pause  at  the  end  of  the  diastole 
in  seconds  to  three  decimals.  Records  secured  under  different  condi- 
tions of  exercise,  etc.,  brought  together  in  a  table  will  usually  show  that  the 
higher  heart-rates  decrease  the  time  of  the  cardiac  cycle  at  the  expense  of 
the  diastole.  In  other  words,  the  time  of  the  systole  remains  fairly 
constant,  while  the  time  of  the  diastole  increases  or  decreases  inversely 
with  the  rate,  a  fact  to  which  Hiirthle  has  drawn  attention,  figure  157. 


THE    FROG  S    HEART 


26l 


3.  The  Rate  and  Sequence  of  the  Contractions  of  the  Frog's  Heart. 
Destroy  the  brain  of  the  frog  and  open  the  thorax,  but  do  not  destroy 
the  pericardium.  Count  the  rate  of  the  heart  per  minute,  then  remove  the 
pericardium  and  make  a  second  determination  after  the  heart  is  exposed 
to  the  air.  The  different  parts  of  the  heart  are  easily  identified  and 
the  contractions  in  definite  sequence  can  be  determined  without  difficulty. 
Make  this  determination  for  the  ventricle,  auricle,  and  sinus  venosus  by 
direct  observation. 

Prepare  a  cardiac  lever  as  shown  in  figure  212  or  216,  taking  special 
care  to  arrange  the  foot  so  that  it  will  not  bind  when  in  motion.  Adjust 
the  foot  of  the  lever  on  the  exposed  ventricle  and  bring  its  point  to  write 


FIG.  212. — Heart  Lever  for  Frog  or  Turtle  Hearts.  This  lever  rests  directly  on 
the  surface  of  the  heart,  the  foot  consisting  of  a  tiny  piece  of  dry  cork  bark.  The 
lever  can  also  be  used  as  in  figure  216,  in  this  case  by  attaching  the  tip  of  the  apex 
of  the  ventricle  by  a  tiny  hook  and  thread  to  the  short  axis  of  the  lever.  Either  de- 
vice can  be  used  for  taking  records  of  cardiac  sequence. 

lightly  on  the  smoked  paper  of  a  recording  cylinder.  This  cylinder  should 
travel  at  the  rate  of  about  2  cm.  per  second  and  its  speed  be  marked  by  the 
writing  point  of  an  electric  magnet.  Take  care  to  adjust  the  time  magnet 
in  a  vertical  line  with  the  writing  point  of  the  heart  lever,  placing  the 
heart  lever  about  i  cm.  above.  The  tracing  of  the  ventricle,  the  cardio- 
gram, will  show  rhythmic  contraction,  relaxation,  and  pause  of  the  ven- 
tricle. It  will  also  enable  one  to  measure  the  exact  proportion  of  the  total 
time  of  the  cardiac  cycle  consumed  by  the  systole  and  diastole,  and  also 
that  portion  of  the  diastole  in  which  the  ventricle  is  wholly  at  rest.  A 
drum  rate  of  2  mm.  per  second  gives  a  more  satisfactory  record  of  varia- 
tions when  amplitude  and  rate  alone  are  studied. 


262 


THE  CIRCULATION  OF  THE  BLOOD 


After  one  has  obtained  ventricular  tracings  and  has  learned  the  diffi- 
culties of  adjusting  the  apparatus,  a  second  heart  lever  should  be  adjusted 
to  the  auricle,  and  the  auricular  movements  recorded  at  the  same  time  as 
those  of  the  ventricle.  If  some  care  is  taken  to  adjust  these  two  writing 
points  in  a  vertical  line  a  splendid  tracing  showing  synchronism  between 


FIG.  213. — Cardiogram  Showing  Contractions  of  the  Auricle,  a,  and  Ventricle,  v 
of  a  Frog.  Time  in  seconds.  The  record  shows  the  sequence  of  the  auricle  and  ventricle, 
(New  figure  by  Dooley.) 

auricle  and  ventricle  is  obtained.  Measure  the  rate  and  the  time  of  the 
different  phases  of  the  contractions  of  the  auricle  and  ventricle  and 
tabulate  them  in  the  following  form,  always  expressing  fractions  in  the 
decimal  system: 


Rate  per 
minute 

Time  of  systole 
in  seconds 

Time  of  diastole 
in  seconds 

Time  of  pause 
in  seconds 

Auricle  

Ventricle 

4.  Sequence,  Conduction  and  Heart  Block,  Turtle  Heart. — Prepare  a 
turtle,  expose  the  heart  and  determine  the  rate  and  sequence  of  the 
parts.  Observe  that  the  veins  are  contractile. 

a.  Make  simultaneous  records  of  the  right  auricle  and  the  ventricle, 
use  speeds  of  2  mm.  and  2  cm.  per  second. 

b.  Attach  a  GaskelPs  clamp  close  against  the  ventricle  in  the  a-v 
groove.     While  taking  records  slowly  compress  the  clamps  by  steps  until 
the  ventricular   rhythm  begins   to   slow  down.     Produce  partial  block 
through  2  auricles  to  i  ventricle,  3  to  i,  4  to  i,  etc.,  rhythms,  until  com- 
plete block  is  obtained.     Note  recovery  on  removing  the  clamp. 

c.  While  recording  the  contractions  of  the  right  and  left  auricles  by 
separate  levers  split  the  heart  into  right  and  left  halves.     The  right  half 
will  continue  the  usual  rhythm,  the  left  will  be  slower  but  show  more 
tone.     These  halves  will  respond  to  nerve  control  and  to  block  tests. 


THE    FROG'S    HEART 


263 


5.  The  Contractions  of  the  Excised  Heart  of  the  Frog — Pith  a  frog 
and  expose  the  heart,  as  described  in  the  preceding  experiment.     Re- 
move it  completely  from  the  body  by  first  cutting  the  arteries  at  their 
branching  in  front  of  the  bulbus  arteriosus,  then  carefully  lifting  up  the 
parts  of  the  heart  and  cutting  away  the  great  veins  where  they  enter  the 
sinus.     This  will  remove  the  entire  heart,  including  all  its  contractile 
parts.     The  frog's  heart,  when  thus  removed  and  still  wet  with  its  own 
blood,  will  continue  contracting  rhythmically  and  in  its  natural  sequence 
for  some  hours.     Place  such  an  isolated  heart  in  a  watch-glass  and  take  a 
record  of  its  contractions  by  the  apparatus  described  in  the  preceding 
experiment.     (The  same  phenomena  may  be  studied  on  a  heart  isolated 
and  mounted  in  a  Williams'  apparatus.) 

Set  this  watch-glass  on  the  metal  warming-box  supplied,  and  arrange 
for  the  circulation  of  water  of  different  temperatures  through  the  box. 
Vary  the  temperature  of  the  box,  and  therefore  of  the  heart  placed  upon  it, 
by  allowing  water  of  o°  C.,  10°  C.,  20°  C.,  30°  C.,  40°  C.,  to  flow  through 
it.  Or  place  the  heart  in  a  watch-glass  over  a  drinking  glass  of  water  of  the 
proper  temperature.  Record  the  contractions  of  the  heart  at  each  of 
these  temperatures.  The  exposed  heart  will  not  take  the  same  absolute 
temperature  as  the  box,  but  the  relative  temperature  will  be  decreased  or 
increased.  Tabulate  the  rates  at  these  different  temperatures  by  the 
plan  previously  described. 

6.  The  Perfused  Heart,  Influence  of  Different  Nutrient  Fluids.— Expose 
a  frog's  heart,  as  previously  described,  and  insert  a  4-way  cannula  into 
the  ascending  vena  cava  where  it 

enters  the  sinus.  Connect  the 
limbs  of  the  venous  cannula  with 
Mariotte's  bottles.  Fill  one  with 
Ringer's  solution,  the  other  with 
comparison  fluids.  Adjust  the  con- 
stant level  tube  for  a  pressure  of 
4  cm.  of  fluid  and  allow  it  to  flow 
through  the  heart.  The  heart  will 
continue  its  contraction's  in  good 
sequence  and  with  a  uniform  rate. 
Record  the  contractions  by  the 
Engelmann  lever  method  on 
smoked  paper,  together  with  a 
time  tracing  in  seconds.  Set  the 
drum  at  the  rate  of  about  2  mm. 

per  second.     After  each  compari- 

-    .  .    .       ..  FIG.  214. — Roy's  Tonometer, 

son  is  made  Ringer  s  fluid  should 

be  perfused  to  secure  a  return  of  the  normal  contractions. 


264  THE    CIRCULATION    OF    THE   BLOOD 

Use  the  tracing  obtained  under  the  influence  of  Ringer's  solution  as  a 
normal  and  compare  it  with  the  rate  and  amplitude  of  the  contractions 
when  the  heart  is  perfused  with: 

a.  Physiological  saline   solution,    then    return    to   Ringer's   solution; 

b.  With  saline  and  potassium  chloride  in  the  proportions  found  in 
Ringer's  solution  (.7  per  cent  sodium  chloride  +  .03  per  cent  potassium 
chloride) ; 

c.  With  saline  and  calcium  chloride  in  the  proportions  found  in  Ringer's 
solution  (.7  per  cent  sodium  chloride  +  .026  per  cent  calcium  chloride);, 

d.  With  Locke's  solution; 

e.  With  milk  diluted  4  volumes  with  physiological  saline; 
/.  With  normal  serum,  or  blood; 

g.  With  blood,  or  serum,  diluted  four  times  with  saline. 
Tabulate  the  rates  and  amplitudes  of  the  heart  under  these  different 
influences  by  the  method  previously  followed. 

7.  The  Heart  Volume. — Isolate  a  frog's  heart  by  the  method  de- 
scribed for  perfusing  it  with  fluid  in  the  preceding  experiment.     Connect 
it  in  a  Roy's  tonometer,  see  figure  214,  adjust  the  lever  of  the  tonometer 
for  a  tracing  on  smoked  paper.     This  instrument  records  the  change  in 
volume  with  each  heart  contraction.     The  influence  of  pressure,  varied 
between  2  and  10  cm.,  and  of  nutrient  fluids  on  the  heart  volume  may  be 
determined. 

An  instructive  demonstration  can  be  had  by  placing  the  heart  of  the 
cat  or  dog  in  a  Henderson  plethysmograph  and  recording  the  volume 
changes  by  the  tambour  method. 

8.  The  Isolated  Heart  of  the  Terrapin.— The  heart  of  the  terrapin, 
being  somewhat  larger  and  somewhat  more  responsive  than  the  heart  of  the 
frog,  may  be  substituted  in  the  two  immediately  preceding  experiments. 
The  facts  obtained  from  it  will  be  essentially  the  same  as  those  obtained 
from  the  frog's  heart. 

9.  The  Isolated  Mammalian  Heart. — The  mammalian  heart  may  be 
isolated  from  the  body  and  kept  alive  and  contracting  for  many  hours, 
as  has  been  demonstrated  by  numerous  observers.     It  is  only  necessary 
to  keep  the  temperature  approximately  that  of  the  normal  body  and  to- 
perfuse  the  heart  through  the  coronary  circulation  with  aerated  blood, 
or  diluted  blood,  containing  sufficient,  aerated  hemoglobin  to  supply  the 
heart  with  the  requisite  amount  of  oxygen.     Or  the  heart  may  be  kept 
alive  on  the  inorganic  salt  solutions,  provided  these  are  supplied  with 
oxygen  under  considerable  tension  (Porter,  Howell).     Even  the  human 
heart  has  been  isolated  and  kept  contracting  for  some  hours  in  the  above 
manner  (Kuliabko).     The  method  used  is  to  insert  a  cannula  into  the 
aorta  and  perfuse  the  heart  through  the  coronary  circulation  under  ade- 
quate pressure,  as  described  by  Martin.     Many  interesting  experiments 


AUTOMATIC    CONTRACTIONS    OF    CARDIAC    MUSCLE  265 

and  demonstrations  can  be  made  on  the  mammalian  heart,  but,  as  this, 
experiment  is  usually  a  demonstration  experiment,  the  detail  of  procedure 
is  left  to  be  supplied  by  the  demonstrator. 

10.  Automatic  Contractions  of  Cardiac  Muscle. — Isolated  portions  of 
the  dog's  ventricle  have  been  kept  in  rhythmic  contraction  by  Porter. 
But  the  best  laboratory  material  is  supplied  by  the  heart  of  the  terrapin. 
Cut  a  strip  from  the  ventricle  of  the  terrapin  extending  around  its  curved 
apex,  as  shown  by  the  dotted  line  in  the  accompanying  figure,  215.  Split 
this  strip  longitudinally  into  two  parts,  each  of  which  will  then  be  about 
3  to  4  mm.  in  diameter  by  3  cm.  long.  Use  care  to  cut  smooth  strips. 
Tie  a  silk  thread  around  the  extreme  tips  of  each  end  of  the  strip,  tying  a 
loop  of  about  i  cm.  long  at  one  end,  and  about  10  cm.  long  at  the  other. 
Suspend  the  strip  over  a  glass  hook,  figure  216,  by  the  short  loop,  and  con- 
nect it  with  a  heart  lever  by  the  long  loop,  as  shown  in  the  same  figure. 
Use  a  tension  of  i  gram.  Contractions  of  this  strip  as  arranged  will  be 
recorded  with  a  magnification  of  about  five  and  with  the  upstroke  of  the 
lever,  which  is  convenient  for  reading  and  interpretation.  Keep  the  strip 
moist  with  physiological  saline  in  a  specimen  tube  i  by  3  inches  in  size. 
The  arrangement  of  apparatus  figured  makes  it  possible  easily  and 
quickly  to  change  this  solution  for  any  other  that  may  be  desired. 


PIG.  215. — Heart  of  the  Terrapin  to  Show  the  Method  of  Cutting  the  Apex  Strip.     Vf 
Ventricle;  Au,  auricles;  Vc,  venae  cavae;  Ao,  aorta. 

Contractions  of  the  ventricular  strip  in  saline  begin  in  from  10  to 
40  minutes  after  the  preparation  is  made,  and  go  through  a  regular  sequence 
of  slight  increase  in  rate  and  amplitude  for  from  10  to  20  minutes,  followed 
by  a  very  constant  rate,  but  gradually  decreasing  amplitude  for  a  period  of 
from  2  to  3  hours,  figure  171.  Saline  is  used  for  the  ventricular  muscle 
because  it  increases  the  rhythmicity  to  a  degree  convenient  for  study. 


266  THE  CIRCULATION  OF  THE  BLOOD 

In  pure  serum  or  blood  the  rhythmicity  is  entirely  absent  though  it  is 
significant  that  the  few  sporadic  contractions  that  do  occur  are  large. 

This  preparation  makes  possible  many  instructive  experiments  tending 
to  show  fundamental  properties  of  cardiac  muscle.  The  preparation 
contains  no  nervous  mechanism  except  terminal  fibers,  and  its  behavior 
may  be  safely  attributed  to  the  muscle  substance  itself. 

Try  the  following  experiments:  i.  Submit  the  strip  to  saline  solutions 
of  different  temperatures,  varying  through  steps  of  10  degrees  from  o°  C. 
to  30°  C.  2.  Try  the  effect  of  the  different  ingredients  in  Ringer's  solution 
using  physiological  saline  as  the  standard  normal  for  the  ventricle,  see  Exp. 
6,  b,  c,  etc. 

a.  Combine  potassium  chloride  with  saline,  figure  172; 

b.  Calcium  chloride  with  saline,  figure  173; 

c.  Potassium  and  calcium  chloride  and  saline; 

d.  Locke's  solution; 

e.  Solution  of  blood  diluted  with  saline; 

/.  Solution  of  milk  with  saline  in  the  proportion  of  one  part  milk  to 
four  of  saline. 

Cut  and  mount  strips  from  the  auricle  and  from  the  sinus,  letting  the 
latter  extend  out  on  to  the  vena  cava.  In  these  last  preparations  care 
must  be  taken  to  balance  the  lever,  as  a  slight  overtension  paralyzes  the 
muscle. 

Immerse  these  strips  in  pure  serum,  or  Ringer  but  not  physiological 
saline,  and  compare  their  behavior  with  that  of  the  ventricle  in  pure  serum. 
The  sinus  and  usually  the  auricle  will  be  found  rhythmic  in  serum,  while 
the  ventricle,  if  it  contracts  at  all,  will  contract  at  irregular  periods. 
Often  there  is  a  distinct  progressive  decrease  in  the  rhythm,  the  sinus 
having  the  same  rhythm  as  the  whole  heart,  the  auricle  a  considerably 
slower  rhythm,  and  the  ventricle  slow  and  aperiodic.  The  sinus  prepa- 
ration will  show  beside  the  fundamental  rhythm  a  characteristic  slow 
contraction  and  relaxation,  which  has  been  described  as  tone,  figure  170. 
Repeat  Exp.  6,  a,  b}  c,  etc.,  on  the  auricle  and  sinus. 

ii.  Influence  of  the  Cardiac  Nerves  on  the  Frog's  Heart. — Care- 
fully pith  a  frog  with  the  minimal  loss  of  the  blood  of  the  animal.  Expose 
the  heart  as  previously  described,  make  a  cut  through  the  manubrium, 
continue  it  through  the  skin  and  muscles  at  the  angle  of  the  jaw,  thus 
exposing  the  vagus  nerve.  The  vagus  runs  along  the  edge  of  a  delicate 
muscle  diagonally  downward  and  backward  toward  the  heart.  The 
glosso-pharyngeal  is  just  in  front  of  the  vagus  and  the  hypoglossal  just 
behind  it.  The  latter  runs  parallel  with  the  vagus  near  its  origin,  but 
lower  down  turns  across  the  vagus  and  runs  to  its  distribution  in  the 
tongue  muscles.  These  two  nerves  serve  to  aid  the  student  in  the  identi- 
fication of  the  vagus,  see  figure  217.  It  is  usually  better  to  cut  the 


INFLUENCE    OF    THE    CARDIAC    NERVES    ON    THE     FROG'S    HEART       267 


hypoglossal  away,  and  also  to  cut  the  brachial  and  the  laryngeal  nerves 
to  prevent  the  stimulation  of  undesired  structures. 


FIG.  216. — Arrangement  of  Apparatus  for  Studying  the  Contractions  of  the  Strip  of  the 

Apex  of  the  Ventricle. 

Prepare  an  induction  coil,  see  laboratory  experiments  on  muscle.  Use 
platinum  electrodes  of  the  Harvard  pattern,  set  the  coil  for  a  mild  stimulus 
tested  by  the  lips  or  the  tongue, 
lift  up  the  vagus  gently  and  lay 
it  on  the  platinum  tips  of  the 
electrodes.  Take  care  that  the 
electrodes  do  not  come  in  contact 
with  the  adjacent  tissue.  Arrange 
a  signal  magnet  as  shown  in  the 
diagram,  so  that  the  magnet  and 
the  stimulating  key  of  the  induc- 
tion coil  may  be  closed  and  opened 
at  the  same  instant.  When  all  is 
ready  (a)  secure  a  normal  record, 
then  (b)  stimulate  the  vagus  for 
five  to  ten  seconds,  recording  the 
time  with  the  signal  magnet  and 
allowing  the  record  to  continue 

until  the  heart  has  returned  to  its 

i  T.     j  FIG.  217. — Diagram   Showing  the  Rela- 

normal  rate  and  amplitude, -I.e.,  tions  of  the f  vago-sympathetic  Nerve  to  the 


Hy,   Hypoglossal;  Gl, 
V,   vago- 


usually   one  circuit  of  the  drum.   Heart,  in  the  Frog. 
Most  students  fail  in  this  experi- 
ment  by  'not   allowing   sufficient 
time  in  the  record  for  a  normal  before  stimulation,  and  by  not  allowing 


268  THE  CIRCULATION  OF  THE  BLOOD 

sufficient  time  after  stimulation  for  a  return  to  the  normal.  It  will  be 
better  to  take  one  good  tracing,  showing  all  the  facts  of  the  experiment, 
than  several  partial  tracings,  none  of  which  are  complete. 

With  these  suggestions  in  mind,  (c)  repeat  the  above  experiment, 
using  stimulating  currents  of  increasing  intensity  until  complete  cardiac 
inhibition  is  produced,  (d)  Perform  experiments  showing  the  influence 
of  the  duration  of  the  stimulation  on  the  inhibition;  i.e.,  stimuli  of  i 
second,  2  seconds,  10  seconds,  and  30  seconds. 

In  the  frog  the  vagus,  or  inhibitory,  and  sympathetic,  or  accelerator, 
fibers,  are  found  in  one  trunk,  the  vago -sympathetic,  but  stimuli  will  usually 
produce  inhibitions  and  not  acceleration.  Occasionally  with  very  weak 
preparations  direct  acceleration  may  be  produced.  To  get  the  pure 
inhibitory  or  pure  accelerator  effects  one  must  dissect  back  to  and  (e) 
stimulate  the  trunk  of  the  vagus  before  it  is  joined  by  the  sympathetic 
fibers;  or  to  the  sympathetic  trunk  and  (/),  stimulate  between  the  third 
spinal  nerve  and  the  point  where  it  joins  the  vagus  trunk.  Pure  accelera- 
tor effects  may  be  demonstrated  by  (g),  stimulating  the  vago-sympathetic 
after  applying  i  c.c.  of  o.i  per  cent,  atropine  to  the  heart  to  poison  the 
vagus  endings,  (ti)  perfusion  of  i  c.c.  of  o.oi  per  cent,  epinephrin  will 
chemically  stimulate  the  accelerator  endings  in  the  presence  of  normal 
vagus  endings. 

In  the  study  of  the  above  experiments  one  should  note  the  rates  per 
minute  and  the  amplitude  of  the  normal  period  just  before  stimulation r 
the  rate  and  amplitude  during  the  period  of  stimulation,  and  the  same 
at  different  times  after  the  stimulation  until  constant  results  are  obtained. 
A  tabulation  of  these  results  will  usually  enable  one  to  judge  the  influence 
of  each  of  the  various  factors  recommended  in  the  experiment. 

12.  Influence  of  the  Cardiac  Nerves  on  the  Terrapin's  Heart.— 
Instead  of  the  frog  one  may  use  the  terrapin  in  the  above  experiment. 
In  this  animal  the  vagus  and  sympathetic  in  the  neck-  can  very  readily  be 
isolated.     It  is  usually  quite  impossible  to  demonstrate  any  cardiac  accel- 
eration.    But  the  vagus  produces  inhibitions  which  differ  from  the  effects 
in  the  frog  in  that  during  the  recovery  from  complete  inhibitions  the 
ventricular  contractions  are  apparently  at  once  maximal,  see  figures  180 
and  181.     In  the  frog  the  ventricular  contractions  when  they  reappear 
are  at  first  slight,  but  gradually  increase  in  amplitude  until  they  have 
their  former  value.     The  student  should  explain  the  significance  of  these 
observations. 

13.  Arterial  Blood-pressure  in  Man. — The  arterial  blood-pressure  in 
man  can  be  measured  indirectly  by  measuring  the  pressure  which  it 
takes  around  the  arm  completely  to  close  the  artery.     Some  form  of  the 
Riva-Rocci  type  of  apparatus,  preferably  the  Tyco  or  Faught,  should  be 
used.     Two  points  fundamental  to  physiology  and  to  clinical  diagnosis  can 
be  determined,  the  systolic  or  maximum  pressure  in  the  artery,  and  the 
diastolic  or  minimum  pressure. 


THE    ARTERIAL   BLOOD-PRESSURE    IN   A    MAMMAL  269 

a.  Bare  the  left  arm,  and  wrap  the  rubber  sleeve  band  snugly  just 
above  the  elbow  and  turn  the  free  end  under  at  the  top.     Connect  the 
Tyco  or  Faught  manometer  with  one  entrance  tube  and  the  bulb  pump 
with  the  other. 

Lightly  bind  the  bell  of  a  stethoscope  over  the  brachial  artery  on  the 
inner  surface  of  the  forearm  just  under  the  border  of  the  arm  band. 

b.  Quickly  fill  the  band  to  a  pressure  of  150  mm.,  i.e.,  completely  com- 
press the  brachial  artery.     Slowly  allow  the  air  to  escape,  listening  care- 
fully with  the  stethoscope  for  the  first  appearance  of  a  pulse  murmur  in 
the  artery.     At  the  same  time  watch  for  the  appearance  of  oscillations 
in  the  dial  of  the  manometer.     At  a  certain  pressure  the  hand  of  the  dial 
will  suddenly  begin  to  oscillate  and  a  distinct  intermittent  sound  clear 
and  sharp  in  tone  will  be  heard.     This  is  the  moment  when  the  first  escape 
of  blood  through  the  compressed  artery  takes  place.     It  measures  the 
systolic  pressure,     c.  Continue  to  reduce  the  pressure.     The  oscillations 
of  the  dial  will  increase  up  to  a  certain  point.     An  intermittent  pulse  will 
be  heard  passing  into  a  loud  and  sharp  snappy  sound.     At  a  certain 
point  the  sound  suddenly  becomes  dull  and  low  and  disappears.     The 
point  of  maximum  oscillation  and  of  disappearance  of  the  intermittent 
sound  marks  the  diastolic  pressure. 

Measure  the  pressures  in  the  standing  position,  in  sitting  position, 
and  after  a  short  run.  Tabulate  the  results  and  draw  averages.  Repeat 
the  measurements  on  children  and  elderly  people,  using  care  not  to  pro- 
long the  total  compression  of  the  artery. 

14.  The  Arterial  Blood-Pressure  in  a  Mammal. — The  arterial  blood- 
pressure  may  be  measured  on  the  anesthetized  cat,  dog,  or  rabbit.  Simple 
blood-pressure  was  originally  measured  by  Hale's  method  of  connecting 
the  artery  with  a  vertical  tube  and  allowing  the  blood  to  flow  freely  into 
the  tube  until  a  column  was  raised  to  the  height  which  balanced  the  pres- 
sure in  the  vessel.  This  simple  method  is  decidedly  the  best  for  the 
beginner,  since  it  does  not  necessitate  the  use  of  very  complicated  appara- 
tus. At  the  same  time  it  gives  practice  in  anesthesia  and  in  operations 
under  anesthetics,  and  therefore  serves  as  a  good  preparation  for  the  more 
complicated  experiments  which  follow. 

The  necessary  apparatus  should  first  be  prepared  as  follows :  A  vertical 
tube  supported  on  a  stand  with  a  scale  graduated  in  the  metric  system, 
assorted  cannulae  of  approximately  the  size  of  the  carotid  artery  of  the  ani- 
mal to  be  operated  on,  linen-thread  ligatures,  dissecting  set  in  good  condition, 
an  animal-holder  with  strings  or  straps  firmly  to  fasten  the  anesthetized 
animal,  a  chloroform-ether  mixture  for  dogs  (or  other  anesthetic  according 
to  the  animal  to  be  used).  Four  men  should  be  assigned  to  perform  this 
experiment.  While  two  are  chloretonizing,  anesthetizing  and  preparing 
the  animal,  two  should  arrange  the  apparatus  as  nearly  ready  for  con- 


270  THE    CIRCULATION    OF    THE    BLOOD 

necting  with  the  artery  as  possible.  When  all  the  apparatus  is  arranged 
and  the  animal  anesthetized,  it  should  be  tied  firmly  to  the  animal-holder. 
Let  one  experimenter  attend  strictly  and  at  all  times  to  anesthetizing  the  animal; 
recovery  to  light  anesthesia  must  not  occur.  Let  the  operator  quickly  expose 
about  3  cm.  of  the  carotid  artery  (or  the  femoral  artery  if  circulation  time 
is  also  to  be  tested  on  this  animal),  by  making  an  incision  through  the  skin 
of  the  neck  5  cm.  long,  and  dissecting  down  between  the  muscles.  Sepa- 
rate the  carotid  from  the  adherent  vagus  nerve  by  tearing  the  connective 
tissue  with  the  scalpel  handle,  freeing  the  vessel  for  about  2  to  3  cm.  of  its 
length.  Lay  two  loose  ligatures  of  linen  thread  around  the  vessel,  place 
a  small  bulldog  forceps  on  the  exposed  artery  nearest  the  heart,  and  ligate 
the  end  nearest  the  head  with  one  of  the  ligatures.  Lift  up  the  inter- 
vening artery  with  strong  forceps  and  make  a  V-shaped  cut  near  the  liga- 
ture pointing  the  cut  toward  the  heart,  let  it  extend  about  half-way  across 
the  artery.  Introduce  a  cannula  toward  the  heart,  and  tie  it  firmly  with 
the  second  ligature.  Connect  the  cannula  with  the  rubber  tubing  to 
the  vertical  glass  tube. 

When  all  is  ready  remove  the  bulldog  forceps  from  the  artery.  The 
blood  will  flow  freely  from  the  artery  into  the  tube,  rising  by  rapid  spurts 
until  the  pressure  from  the  column  of  liquid  is  just  equal  to  that  inside 
the  artery  itself.  If  an  anticoagulant-like  powdered  potassium  oxalate 
is  first  introduced  into  the  vertical  tube,  probably  clotting  at  the  cannula 
will  be  delayed  for  some  minutes.  The  mounting  of  the  blood  into  the 
empty  tube  makes,  indeed,  a  most  striking  demonstration. 

An  accurate  measure  of  the  height  of  the  top  of  the  column  above  the 
level  of  the  cannula  at  the  artery  represents  the  arterial  blood-pressure  in 
terms  of  blood.  The  specific  gravity  of  blood  is  1.056;  of  mercury,  13.6. 
Record  the  pressure  you  obtain  in  terms  of  blood  and  of  mercury.  Note 
also  the  variations  in  pressure  and  account  for  the  rhythm  of  each. 
There  will  be  a  general  variation  of  pressure,  depending  upon  the  degree 
of  anesthesia.  If  anesthesia  is  light  and  muscular  movements  happen, 
there  will  be  an  increase  in  the  blood-pressure.  If  the  anesthesia  is 
heavy,  then  the  blood-pressure  falls.  These  points  of  variation  should 
be  marked  and  recorded  at  once  in  note-books.  Make  full  notes  of  all 
accessory  facts  which  would  aid  in  explaining  the  variations  in  blood- 
pressure,  such  as  size  of  the  animal,  rate  of  respiration,  rate  of  heart-beat, 
the  variations  in  anesthesia,  the  presence  of  the  reflexes,  etc.,  etc. 

Chloroform  the  animal  to  kill  it,  and  note  the  change  in  blood-pressure 
during  the  process,  but  first  do  experiment  15. 

15.  The  Circulation  Time. — The  circulation  time  is  most  satisfactorily 
determined  in  the  laboratory  by  introducing  a  saline  solution  of  methylene 
blue  into  the  jugular  vein  on  one  side.  Note  the  exact  time  with  a  stop- 
watch until  the  color  appears  in  the  carotid  artery,  and  in  the  jugular 
vein  of  the  opposite  side. 


THE   BLOOD-PRESSURE    MODEL  271 

Anesthetize  a  cat  or  dog  with  a  chloroform-ether  mixture,  tie  it  on  the 
animal-holder  and,  when  the  eye  reflexes  are  lost,  expose  the  jugular  vein 
on  the  right  side,  the  carotid  artery  and  the  jugular  vein  on  the  left.  Fill  a 
2-cm.  hypodermic  syringe  with  i  per  cent,  methylene  blue  in  physiological 
saline,  insert  the  needle  into  the  right  jugular  vein,  pointing  it  toward  the 
heart.  Lift  the  left  carotid  artery  and  place  under  it  a  strip  of  moist  white 
paper  2  cm.  wide;  prepare  the  left  jugular  vein  in  the  same  way.  Place  the 
animal  so  that  these  vessels  are  lighted  to  the  best  advantage.  At  a  given 
moment  inject  the  contents  of  the  hypodermic  syringe,  noting  the  time 
with  a  stop-watch.  Observe  the  color  of  the  left  carotid  and  the  left 
jugular,  respectively,  very  carefully,  and  take  the  time  of  the  first  appear- 
ance of  the  methylene  blue.  The  color  will  appear  first  in  the  artery, 
second  in  the  vein.  The  difference  in  time  between  the  moment  of  injec- 
tion and  the  moment  of  color  in  the  artery  represents,  with  a  slight  cor- 
rection, the  circulation  time  of  the  pulmonary  or  lesser  circulation.  The 
time  from  the  injection  until  the  color  in  the  other  jugular  vein  represents 
the  total  time  of  circulation. 

Stewart  has  made  these  determinations  even  more  correctly  by  the  elec- 
trical-resistance method.  He  injected  10  per  cent,  salt  solution  and  deter- 
mined the  variation  in  resistance  by  a  galvanometer.  If  the  galvanometer 
is  available,  then  check  the  above  determinations  by  the  electrical  method, 
arranging  the  apparatus  under  the  direction  of  an  instructor. 

1 6.  The  Blood -pressure  Model. — An  artificial  model  of  the  circulatory 
apparatus,  which  illustrates  all  mechanical  parts  involved,  has  been 
arranged  by  Porter,  or  can  be  easily  constructed.  The  model  should 
have  the  following  possibilities:  A  pump,  which  permits  of  rhythmic 
action  at  a  varying  rate  and  varying  force;  a  resistance  to  the  outflow 
liquid  which  can  be  increased  or  decreased;  and  an  elastic  set  of  vessels 
into  which  the  pump  discharges. 

If  Porter's  schema  is  used,  determine  the  following  points,  (a)  The 
pressure  in  terms  of  mercury  in  the  arterial  and  venous  limbs  of  the 
apparatus  when  the  pump  makes  a  rate  of  72  per  minute;  (b)  the  influence 
on  these  two  pressures  when  the  rate  is  increased,  when  it  is  decreased, 
(c)  the  effect  on  these  pressures  when  the  peripheral  resistance  is  great, 
when  it  is  low.  With  a  sphygmograph,  (d)  take  a  tracing  of  the  pulse  in 
the  elastic  tube  representing  the  arterial  side  of  the  schema. 

If  an  ordinary  bulb  syringe  and  simple  apparatus  is  used,  then  deter- 
mine the  following:  (e)  The  character  and  rate  of  the  outflow  when  water 
is  pumped  into  the  rigid  glass  tube  with  no  resistance  to  the  outflow;  (f) 
when  a  glass  tube  of  smaller  caliber  is  connected  with  the  end  of  the  larger 
glass  tube  so  as  to  produce  a  high  resistance  to  the  outflow,  (g)  Pump  the 
water  into  a  rubber  tube  of  smaller  size  and  compare  with  the  proceeding 
when  there  is  no  resistance  to  the  outflow;  also  (/?)  when  a  glass  tube  of 


272  THE    CIRCULATION    OF    THE    BLOOD 

small  caliber  is  introduced  into  the  end  in  order  to  produce  high  resistance 
to  the  outflow.  Determine  the  amount  of  resistance  necessary  to  produce 
a  constant  outflow  when  the  pump  has  a  rate  of  72  beats  per  minute. 

In  this  experiment  what  effect  is  produced  on  the  outflow  if  the  rate 
of  the  pump  is  varied?  if  the  force  of  the  stroke  is  varied?  if  the  elasticity 
of  the  rubber  tube  representing  the  artery  is  varied?  If  the  resistance 
represented  by  the  size  of  the  glass  tube  at  the  outflow  is  varied? 

17.  The  Arterial  Pulse. — The  form  of  the  arterial  pulse  maybe  taken 
by  one  of  the  various  sphygmographs  applied  to  the  radial  artery  at  the 
wrist  or  the  common  carotid  in  the  neck.     If  the  tambour  method  is  used, 
apply  a  sphygmographic  tambour  on  the  wrist  adjusting  the  central 
pressure  over  the  radial  artery.     Fasten  it  in  place  by  the  proper  bands, 
adjusting  the  tension  by  flexing  the  wrist.     Connect  the  receiving  tambour 
with  a  delicately  balanced,  small-sized  recording  tambour,  which  should 
write  its  movements  on  a  cylinder  revolving  at  the  rate  of  i  to  2  cm.  per 
second. 

A  convenient  clinical  instrument  is  the  Dudgeon  or  the  Jacquet 
sphygmograph.  These  are  to  be  applied  at  the  wrist  and  give  tracings 
showing  delicate  variations  in  the  form  of  the  pulse  wave  with  great 
magnification  and  a  considerable  degree  of  accuracy.  Make  a  comparison 
of  the  form  of  the  pulse  wave  from  tracings  taken  from  at  least  six  different 
individuals. 

The  sphygmogram  from  the  carotid  artery  may  best  be  taken  by 
applying  an  air  sphygmograph  to  the  neck  over  the  carotid  and  fastening 
it  in  position  by  a  spring.  Record  by  the  tambour  method. 

1 8.  The  Rate  of  Propagation  of  the  Pulse  Wave. — Apply  tambour 
sphygmographs  to  the  carotid  in  the  neck  and  to  the  radial  at  the  wrist,  and 
make   simultaneous   records   on   a   drum,   adjusting   the   writing  levers 
of  the  two  recording  tambours  in  an  exact  vertical  line.     Let  the  recording 
drum  travel  at  the  speed  of  2  cm.  or  more  per  second,  and  record  the  speed 
by  a  50  double-vibration  tuning-fork.     The  carotid  pulse  will  be  found  to 
precede  the  radial  pulse  by  the  fraction  of  a  second.     This  short  interval, 
which  can  be  determined  in  thousandths  of  a  second  by  comparison  with 
the  time  tracing  below,  represents  the  time  required  for  the  pulse  wave  to 
traverse  the  distance  from  the  carotid  to  the  radial.     Measure  the  distance 
used  in  the  experiment  and  calculate  the  rate  of  propagation  in  centi- 
meters per  second. 

If  the  writing  points  of  the  recording  levers  in  this  experiment  are  made 
of  very  delicate  strips  of  note  paper  or  of  thin  photographic  film  celluloid, 
so  as  to  offer  little  resistance  to  the  surface  of  the  drum,  the  detail  of  the 
pulse  wave  at  the  two  points  will  be  accurately  transcribed  and  may  be 
compared. 

19.  The    Capillary    Circulation.— The    capillary    circulation  is  best 


CAPILLARY   BLOOD-PRESSURE    OF    MAN  273 

demonstrated  in  the  laboratory  by  direct  observation  on  the  web  of  the 
frog's  foot  by  the  use  of  the  compound  microscope.  Give  a  4o-gm.  frog  a 
hypodermic  injection  of  0.3  c.c.  of  ether  under  the  skin  of  the  back.  Wet 
a  piece  of  cheesecloth  the  size  of  a  handkerchief  with  tap  water  and  wrap 
the  etherized  frog  so  as  to  cover  the  entire  body  with  the  exception  of  the 
foot.  When  the  anesthesia  has  progressed  so  as  to  destroy  voluntary 
movements,  bind  the  foot  on  an  ordinary  frog  board  and  spread  the  web 
over  the  window  in  the  board.  Choose  an  area  of  the  skin  which  shows 
small  arteries,  capillaries,  and  veins,  and  in  which  the  blood  is  flowing 
freely  and  rapidly.  Examine  with  the  low-power  of  a  compound  micro- 
scope. In  a  favorable  field  small  arteries,  capillaries,  and  veins  with 
blood  flowing  rapidly  through  them  will  be  easily  found.  Choose  one 
such  field,  cover  with  a  piece  of  thin  cover-glass,  moisten  with  a  drop 
of  water  if  necessary,  and  examine  with  a  high  power.  Note  in  the  small 
artery  the  pulsating  current;  the  border  of  clear  fluid  along  the  side  of  the 
main  stream  of  blood;  the  white  corpuscles  along  the  clear  borders  of  the 
current.  In  the  small  veins  there  are  usually  no  pulsations  and  the  speed 
of  the  current  is  somewhat  less.  Careful  examination  of  the  capillaries 
will  reveal  a  delicate  wall,  the  individual  corpuscles,  and  the  fact  that  the 
red  corpuscles  are  actually  larger  than  the  diameter  of  the  capillary  at 
some  points  and  must  be  bent  to  pass  through.  Note  that  the  capillaries 
form  an  intricate  and  anastomosing  network;  that  the  current  may  occa- 
sionally be  reversed  in  some  of  the  anastomoses. 

The  anesthetizing  effect  of  the  ether  recommended  will  usually  con- 
tinue about  10  to  20  minutes.  If  the  observation  is  more  prolonged,  a 
second  dose  of  ether  should  be  given.  The  capillaries  in  the  tails  of  small 
fish  are  often  very  readily  observed,  and  these  may  be  substituted  for  the 
frog's  web. 

20.  Capillary  Blood-Pressure  of  Man. — Measure  the  capillary  blood- 
pressure  in  your  own  finger  by  von  Krie's  method.  This  apparatus  con- 
sists of  a  small  piece  of  glass  an  inch  square,  or  less,  which  is  placed  across 
the  knuckle  of  the  finger  just  back  of  the  nail.  A  small  weight  pan  is 
suspended  by  a  loop  of  thread  over  this  glass  plate  so  that  weights  put  in 
the  pan  will  bring  varying  pressure  on  the  plate  above.  Add  weights  to  the 
pan  until  an  area  of  the  skin,  about  5  mm.  in  diameter,  is  blanched  by  the 
pressure.  Mark  the  outline  of  this  bloodless  area  on  the  glass,  take 
off  the  apparatus  and  measure  the  exact  area  of  the  glass  so  marked,  weigh 
the  entire  apparatus  and  compute  the  pressure  in  grams  per  square  centi- 
meter for  the  area.  This  pressure  in  terms  of  mercury  represents  the 
capillary  blood-pressure  in  the  vessels  of  the  skin  of  the  finger  at  that 
level.  Measure  the  pressure  when  the  finger  is  held  at  the  level  of  the  top 
of  the  head;  with  the  finger  held  as  low  as  possible;  held  at  the  level  of  the 
heart.  Tabulate  the  measurements.  The  capillary  blood-pressure  at  the 
level  of  the  heart  is  usually  from  40  to  50  mm.  of  mercury. 


274  THE    CIRCULATION    OF    THE   BLOOD 

Lombard  determines  the  end  point  in  this  experiment  more  accurately 
by  the  aid  of  a  low  power  binocular  microscope  after  rubbing  vaseline  into 
the  skin  to  increase  its  transparency.  Hooker's  method  of  compressing 
the  capillaries  by  an  air  system  controlled  and  measured  by  a  manometer 
allows  measurement  both  during  compression  and  in  decompression.  This 
method  has  been  adapted  to  the  mammalian  mesentery  by  Ellis. 

21.  The  Arterial  Blood -Pressure  in  a  Mammal  and  Its  Nervous 
Regulation. — After  the  student  has  measured  the  arterial  blood-pres- 
sure by  Hale's  method,  described  above,  he  is  in  a  position  to  study  the 
variations  and  co-ordinations  in  the  blood  circulatory  apparatus.  The  re- 
cording apparatus  consists  of  writing  pens,  seconds  time  marker,  signal 
marker,  blood-pressure  manometer  preferably  Ludwig's  mercury  manom- 
eter, and  a  continuous  paper  kymograph  preferably  Ludwig's  weight- 
driven  form  or  the  Harvard  belt  kymograph  for  a  continuous  record  of  the 
arterial  blood-pressure.  Connect  the  cannula  with  the  mercury  man- 
ometer which  is  provided  with  a  pressure  bottle.  Use  a  cannula  of  the 
form  shown  in  figure  185,  connecting  the  side  limb  of  the  cannula  with  the 
mercury  manometer,  and  the  end  limb  with  the  pressure  bottle.  When 
the  apparatus  is  ready  chloretonize  and  anesthetize  a  mammal  (dog,  cat,  or 
rabbit),  and  bind  it  to  the  animal-holder.  Let  one  operator  attend  strictly 
and  at  all  times  to  the  anesthetic,  for  the  animal  must  not  under  any  condition 
recover  consciousness  during  the  experiment. 

Expose  the  carotid  artery  in  the  neck,  as  described  in  Experiment  13 
above,  arrange  it  with  ligatures  for  inserting  the  cannula,  expose  the 
vagus  nerve  with  the  same  care,  and  throw  ligatures  around  it  for  con- 
venience in  lifting  it  out  of  its  bed.  Make  a  V-shaped  cut  in  the  carotid 
directed  toward  the  heart,  insert  and  ligate  the  cannula  as  previously 
described.  Before  beginning  the  experiment  one  should  see  that  all  the 
tubes  are  filled  with  the  anticoagulating  liquid  and  that  the  manometer  is 
under  pressure  from  130  to  140  mm.  mercury.  When  all  is  ready  start  the 
kymograph,  see  that  the  recording  pens  are  properly  adjusted,  remove  the 
bulldog  forceps  from  the  artery,  and  the  pressure  record  will  begin. 

a.  Take  a  tracing  of  the  normal  arterial  pressure  and  heart  rhythm  with 
the  recording  paper  moving  at  the  rate  of  0.5  cm.  per  second. 

b.  Stimulate  the  right  vagus  nerve  with  a  mild  induction  current  for 
10  seconds.     If  this  stimulus  is  strong  enough  to  produce  change  in  blood- 
pressure  or  inhibitions  of  the  heart-rate,  then  allow  sufficient  time  follow- 
ing the  stimulus  for  the  blood-pressure  to  return  to  the  previous  normal. 
Observing  these  rules,  vary  the  intensity  of  the  stimulus  from  that  which 
produces  no  apparent  effect  to  that  which  produces  complete  inhibition  of 
the  heart.     Vary  the  time  of  the  stimulus  through  i,  5,  io,and  20  seconds, 
using  different  strengths.     Do  not  allow  the  nerve  to  cool,  become  dry, 
or  to  be  unduly  stretched. 


THE    ARTERIAL   BLOOD-PRESSURE  275 

c.  Test  the  sensitiveness  of  the  left  vagus. 

d.  Allow  the  vagus  to  fall  back  in  its  warm  bed  and  stimulate  the  skin 
of  the  animal  at  some  sensory  surface,  say  the  lips,  the  ear,  or  the  foot. 
By  varying  the  intensity  of  the  stimulus,  a  strength  will  be  found  which 
will  produce  no  reflexes  of  the  voluntary  muscles,  but  will  produce  marked 
effects  on  the  heart  rate  and  on  the  blood-pressure. 

Expose  the  sciatic  nerve,  or  any  other  nerve  trunk  containing  afferent 
or  sensory  fibers,  cut  it,  and  stimulate  the  central  end  for  five  seconds. 
With  a  proper  strength  of  stimulus  a  greater  effect  is  produced  on  the 
heart  and  on  the  blood-pressure  than  by  stimulating  a  small  spot  of  skin. 
This  stimulus  will  also  reflexly  accelerate  respiration. 

e.  Cut  the  right  vagus  nerve  and  mark  the  exact  time  on  the  tracing  by 
the  signal  marker.     Do  not  disturb  the  animal  of  the  record  until  stable 
equilibrium  is  again  reached. 

/.  Now  lift  up  the  distal  end  of  the  divided  right  vagus,  and  stimulate 
it  with  the  strength  which  previously  just  produced  inhibition.  Repeat 
the  experiment  on  the  proximal  end  of  the  divided  vagus.  The  reflex 
effects  are  still  threefold,  cardiac,  vasomotor  and  respiratory.  The  stimu- 
lation of  the  proximal  end  of  the  vagus  produces  effects  on  the  heart  rate 
when  one  vagus  is  still  intact.  See  Experiment  /  below. 

g.  After  10  to  15  seconds  cut  the  left  vagus,  marking  the  time  of  cutting 
on  the  tracing  with  the  same  care  as  before.  As  soon  as  the  vagus  nerves 
are  cut,  the  heart-rate  will  be  observed  to  increase  sharply  and  the  blood- 
pressure  to  rise.  The  respirations  also  change  in  rate  and  depth,  a  fact 
which  can  be  noted  directly  and  by  its  influence  on  the  blood-pressure 
tracing. 

h.  Lift  up  the  distal  end  of  the  left  vagus,  and  stimulate  it  with  an 
electric  current  of  the  strength  which  previously  just  produced  inhibition. 
Stimulate  the  proximal  end  of  the  divided  vagus.  The  stimulation 
produces  no  direct  inhibitory  effect  on  the  heart  rate  when  both  vagi  are 
cut,  but  does  produce  profound  changes  in  the  blood-pressure  owing  to 
vaso-motor  reflexes. 

Occasionally  an  animal  will  be  found  in  which  one  or  both  vagi  are 
comparatively  inactive. 

k.  If  the  rabbit  is  used,  stimulate  the  depressor  nerve,  which  produces 
a  marked  fall  in  blood-pressure  from  reflex  effects,  explain. 

/.  Repeat  the  stimulation  of  the  central  end  of  the  sciatic  as  described 
in  /,  now  that  the  vagus  nerves  are  cut.  The  stimulation  of  this  nerve 
no  longer  produces  decrease  in  the  heart-rate,  but  occasionally  an  accelera- 
tion. The  blood-pressure  is  influenced  as  before,  showing  that  the  vaso- 
motor centers  are  reflexly  stimulated. 

m.  When  you  have  finished  the  outline  of  experiments,  give  an  excess 
of  ether  to  kill  the  animal  and  continue  the  record  until  it  is  dead.  The 


276  THE    CIRCULATION    OF    THE    BLOOD 

blood-pressure  will  fall  rapidly,  the  heart-rate  will  become  slower  but 
does  not  cease  for  a  long  time  after  respirations  stop. 

Should  a  clot  form  in  the  cannula,  put  a  bulldog  forceps  on  the  artery, 
disconnect  the  manometer  tube,  and  wash  the  clot  out  by  a  stream  of 
liquid  from  the  pressure  bottle.  Use  care  not  to  allow  this  fluid  to  enter 
the  exposed  wound. 

Represent  the  results  of  each  individual  experiment  in  the  above  series 
in  tabulated  form  which  shall  show:  i,  the  blood-pressure  and  heart-rate 
just  before  each  experiment;  2,  during  the  experiment;  and  3,  at  different 
times  after  the  experiment  until  the  normal  is  reached.  After  the  data 
are  taken  from  the  tracings  and  arranged  in  tabular  form,  make  a  study 
of  these  facts  and  draw  all  the  conclusions  you  can  concerning  the  nervous 
regulations  of  the  heart  and  of  the  blood-pressure.  Make  a  written  report. 

2  2 .  The  Vaso-motor  Changes  in  the  Finger,  the  Plethysmogram.—  Insert 
the  ringer  in  the  Porter  ringer  plethysmograph,  fill  the  tube  with  warmed 
water,  and  connect  it  with  a  small-sized  air  tambour.  The  variations  in 
volume  of  the  finger  are  slight,  so  that  one  must  use  a  most  delicate 
recorder.  Take  a  tracing  at  a  slow  speed,  i  mm.  per  second.  The  finger 
and  its  plethysmograph  should  be  suspended  so  that  no  mechanical 
movements  will  destroy  the  accuracy  of  the  record.  Observations  through 
several  minutes  will  usually  show  variations  in  volume  of  the  finger, 
which  will  be  recorded  by  the  tambour.  The  reagent  must  be  warm 
and  relaxed. 

Try  a  short  mental  problem.  Cold  air  or  cold  water  in  the  face  will 
usually  be  marked  by  a  decrease  in  volume  indicating  vaso-constriction. 
Warmth  will  lead  to  increase  in  volume  indicating  vaso-dilatation.  In 
sleep  there  is  the  greatest  relaxation  and  a  large  volume  pulse  will  be 
present. 

23.  The  Vaso-motors  of  the  Frog's  Web. — Prepare  a  frog  for  observa- 
tion of  the  circulation  of  the  web  under  the  microscope,  as  described  above, 
giving  it  3  drops  of  ether,  or  just  enough  i  per  cent,  curare  to  destroy 
voluntary  movements.  Quickly  dissect  the  sciatic  nerve  in  the  thigh, 
using  extreme  care  not  to  interfere  with  the  circulation.  Mount  the 
preparation,  pick  out  an  active  field  of  capillaries,  small  arteries,  and 
veins  under  the  low  power  of  the  microscope,  then  adjust  the  high  power 
to  a  field  which  shows  one  or  more  small  arteries.  Make  a  drawing  to 
record  the  diameter  of  these  arteries,  using  pigment  cells  for  land-marks, 
or  measure  with  an  ocular  micrometer.  Now  quickly  stimulate  the 
exposed  sciatic  nerve  while  keeping  the  selected  artery  under  constant 
observation.  After  stimulation  for  10  seconds  the  diameter  of  the  vessels 
will  be  seen  to  decrease  considerably,  sometimes  to  the  point  of  complete 
occlusion.  When  the  stimulation  ceases,  the  vessel  will  remain  contracted 
for  a  few  seconds,  then  will  slowly  regain  its  usual  caliber,  figure  201. 


THE    PLETHYSMOGRAM    OF    THE    KIDNEY  277 

This  is  an  exceptionally  good  method  for  direct  observation  of  the  vaso- 
motor  changes. 

24.  The  Plethysmogram  of  the  Kidney. — Anesthetize  a  dog  or  cat, 
see  Experiments  12  and  19  above,  and  take  continuous  blood-pressure 
tracings.  Now  open  the  abdominal  wall  by  an  incision  along  the  median 
line,  expose  the  left  kidney  and  carefully  dissect  off  its  capsule,  taking 
care  not  to  injure  its  artery  and  vein.  Enclose  the  kidney  in  the  renal 
onkometer  and  carefully  seal  with  vaseline  and  cover  with  omentum. 
Connect  it  with  a  delicate  volume  recording  apparatus.  Brodie's  bellows 
recorder  or  a  large  air  tambour  is  the  best  for  this  purpose.  Adjust  the 
recording  apparatus  in  the  vertical  line  with  the  manometer  and  signal 
pens. 

Stimulation  of  the  nerves  which  affect  general  blood-pressure  through 
the  medium  of  the  heart  will  produce  changes  in  the  volume  of  the  kidney 
in  the  same  direction  as  the  blood-pressure.  Stimuli  which  give  varia- 
tions of  the  blood-pressure  without  direct  change  in  the  heart  itself  affect 
the  volume  of  the  kidney  independent  of  the  blood-pressure : 

a.  Dissect  out  and  stimulate  the  splanchnic  nerves  just  where  they 
pass  through  the  pillars  of  the  diaphragm.     They  cause  vaso-constriction 
in  the  kidney  without  sharply  affecting  the  blood-pressure. 

b.  Stimulate  the  depressor  nerve,  or  the  central  end  of  the  divided 
vagus.     The  volume  of  the  kidney  will  increase  though  the  general  blood- 
pressure  decreases,   showing  that  the  fall  of  blood-pressure  is  due  to 
peripheral  vascular  dilatation. 

c.  Stimulate  the  peripheral  end  of  the  divided  vagus  so  as  to  slow  or 
even  completely  to  stop  the  heart.     The  sharp  fall  in  blood-pressure  is  now 
accompanied  by  decrease  in  the  volume  of  the  kidney,  showing  that  the 
kidney  volume  is  merely  passively  following  the  blood-pressure. 


CHAPTER  VI 
RESPIRATION 

THE  maintenance  of  animal  life  necessitates  the  continual  absorption  of 
oxygen  and  the  excretion  of  carbon  dioxide  by  the  living  tissues.  The  blood 
is  the  medium  in  all  animals  which  possess  a  well-developed  blood-vascular 
system  by  which  these  gases  are  carried.  Oxygen  is  absorbed  by  the  blood 
from  without  and  conveyed  to  all  parts  of  the  organism;  and  carbon  dioxide 
which  comes  from  the  cells  within  is  carried  by  the  blood  to  the  surfaces 
from  which  it  may  escape  from  the  body.  The  two  processes — absorption 
of  oxygen  and  excretion  of  carbon  dioxide — are  complementary,  and  their 
sum  is  termed  the  process  of  Respiration. 

In  all  Vertebrata  and  in  a  large  number  of  Invertebrata  certain  parts, 
either  lungs  or  gills t  are  especially  constructed  for  bringing  the  blood  into 
proximity  with  the  aerating  medium  (atmospheric  air,  or  water  containing 
air  in  solution).  In  some  of  the  lower  Vertebrata  (frogs  and  other  naked 
Amphibia)  the  skin  is  important  as  a  respiratory  organ,  and  is  capable  of 
supplementing  to  some  extent  the  functions'  of  the  proper  breathing 
apparatus. 

A  lung  or  a  gill  is  constructed  essentially  of  a  fine  transparent  membrane, 
one  surface  of  which  is  exposed  to  the  air  or  water,  as  the  case  may  be,  while 
on  the  other  surface  is  a  network  of  blood  vessels.  The  only  separation  be- 
tween the  blood  and  aerating  medium  is  the  thin  wall  of  the  blood  vessels 
and  the  thin  membrane  on  which  the  vessels  are  distributed.  The  difference 
between  the  simplest  and  the  most  complicated  respiratory  membrane  is 
one  of  degree  only. 

In  the  mammals  and  the  higher  vertebrates  the  respiratory  membrane 
is  included  within  a  respiratory  cavity,  the  chest  or  thorax,  which  carries  on 
regular  movements,  the  respiratory  movements,  to  bring  changes  of  air  into 
close  contact  with  the  respiratory  surface. 

The  complexity  of  the  respiratory  membrane,  the  kind  of  aerating  me- 
dium, and  the  respiratory  movements  are  not,  however,  the  only  conditions 
which  cause  a  difference  in  the  respiratory  capacity  of  different  animals. 
The  quantity  and  composition  of  the  blood,  especially  as  regards  the  number 
and  size  of  the  red  corpuscles,  and  the  vigor  and  efficiency  of  the  circulatory 
apparatus  in  driving  the  blood  to  and  fro  between  the  lungs  and  the  active 
tissues,  these  are  conditions  of  equal,  if  not  greater,  importance. 

It  may  be  as  well  to  state  here  that  the  lungs  are  only  the  medium  for  the 
exchange,  on  the  part  of  the  blood,  of  carbon  dioxide  for  oxygen.  The 

278 


THE    RESPIRATORY    APPARATUS 


279 


living  tissues  are  the  seat  of  those  combustion  processes  which  consume 
oxygen  and  produce  carbon  dioxide.  These  processes  occur  in  all  parts  of 
the  body  in  the  substance  of  the  living  active  tissues,  and  are  the  true  respira- 
tory processes,  sometimes  called  internal  or  tissue  respiration. 


THE  RESPIRATORY  APPARATUS. 

The  object  of  the  respiratory  movements  being  the  interchange  of  gases 
in  the  lungs,  it  is  necessary  that  the  atmospheric  air  shall  pass  into  them 
and  that  the  changed  air  shall  be  ex- 
pelled from  them.  The  lungs  are 
contained  in  the  chest  or  thorax,  which 
is  a  closed  cavity  having  no  communi- 
cation with  the  outside  except  by 
means  of  the  respiratory  passages. 
The  air  enters  these  passages  through 
the  nostrils  or  through  the  mouth, 
thence  it  passes  through  the  larynx 
into  the  trachea  or  windpipe,  which 
about  the  middle  of  the  chest  divides 
into  two  tubes,  the  bronchi,  one  to 
each  lung. 

The  Larynx.  —  The  upper  part  of 
the  passage  which  leads  exclusively 
to  the  lung  is  formed  by  the  thyroid, 
cricoid,  and  arytenoid  cartilages, 
figure  218,  and  contains  the  vocal  cords, 
by  the  vibration  of  which  the  voice  is 
chiefly  produced.  These  vocal  cords 
are  ligamentous  bands  covered  with 
mucous  membrane  and  attached  to 
certain  cartilages  which  are  capable  of 
movement  by  muscles.  By  their  ap- 
proximation the  cords  can  entirely 
close  the  entrance  into  the  larynx;  but 
under  ordinary  conditions  the  entrance 
of  the  larynx  is  formed  by  a  more  or 
less  triangular  opening  between  them, 
called  the  rima  glottidis.  Projecting 
at  an  acute  angle  between  the  base  of 
the  tongue  and  the  larynx  to  which  it 

is   attached,  is  a  leaf-  shaped   cartilage     the  trachea,  showing  sixteen  cartilaginous 
.        -  ^.  .       rings;  b,  the  right,  and  b',  the  left  bronchus. 


FIG.  218. — Outline  Showing  the  General 
Form  of  the  Larynx,  Trachea,  and  Bronchi, 
as  seen  from  Before,  h,  The  great  cornu 
of  the  hyoid  bone;  e,  epiglottis;  /,  superior, 
and  tr,  inferior  cornu  of  the  thyroid  carti- 
lage; c,  middle  of  the  cricoid  cartilage;  tr, 


.,,._,,  .        -  ^.  . 

with  its  larger  extremity  free.      This 


(Allen  Thomson.) 


280 


RESPIRATION 


is  called  the  epiglottis.  The  whole  of  the  larynx  is  lined  by  mucous  mem- 
brane, which,  however,  is  extremely  thin  over  the  vocal  cords.  At  its  lower 
extremity  the  larynx  joins  the  trachea. 

Taste  buds  have  been  found  in  the  epithelium  of  the  posterior  surface  of 

the  epiglottis,  and  in  several  other 
situations  in  the  laryngeal  mucous 
membrane. 

The  Trachea  and  Bronchi.— 
The  trachea  extends  from  the 
cricoid  cartilage,  which  is  on  a 
level  with  the  fifth  cervical  vertebra, 
to  a  point  opposite  the  third  dorsal 
vertebra,  where  it  divides  into  the 
two  bronchi,  one  for  each  lung, 
figure  218.  The  trachea  measures, 
on  an  average,  four  or  four  and  a 
half  inches,  12  to  14  cm.,  in  length, 
and  from  three-quarters  of  an  inch 
to  an  inch,  2  to  2.5  cm.,  in  diameter, 
and  is  essentially  a  tube  of  fibro- 
elastic  membrane  within  the  layers 
of  which  are  enclosed  a  series  of 
cartilaginous  rings,  from  sixteen  to 
twenty  in  number.  These  rings 
extend  only  around  the  front  and 
sides  of  the  trachea,  about  two- 
thirds  of  its  circumference,  and 
are  deficient  behind;  the  interval 
between  their  posterior  extremities 
being  bridged  over  by  a  continua- 
tion of  the  fibrous  membrane  in 
which  they  are  enclosed,  figure 
219,  h. 

Immediately  within  this  tube  and 


FIG.  219. — Section  of  the  Trachea,  a, 
Columnar  ciliated  epithelium;  6,  and  c, 
proper  structure  of  the  mucous  membrane, 
containing  elastic  fibers  cut  across  trans- 
versely; d,  submucous  tissue  containing 
mucous  glands,  e,  separated  from  the  hya- 
line cartilage,  g,  by  a  fine  fibrous  tissue,/,  h, 
external  investment  of  fine  fibrous  tissue. 
(S.  K.  Alcock.) 


at  the  back  is  a  layer  of  unstriped 
muscular  fibers.  This  muscular  layer  extends  transversely  between  the 
ends  of  the  cartilaginous  rings  to  which  it  is  attached,  and  also  opposite  the 
intervals  between  them;  its  evident  function  being  to  diminish  the  caliber 
of  the  trachea  by  approximating  the  ends  of  the  cartilages.  Outside  there 
are  a  few  longitudinal  bundles  of  muscular  tissue,  which,  like  the  preceding, 
are  attached  both  to  the  fibrous  and  to  the  cartilaginous  framework. 

The  mucous  membrane,  figures  219  and  220,  consists  largely  of  adenoid 
tissue,  separated  from  the  stratified  columnar  epithelium ,  which  lines  it,  by  a 


THE  TRACHEA  AND  BRONCHI 


28l 


homogeneous  basement  membrane.  This  is  penetrated  here  and  there  by- 
channels  which  connect  the  adenoid  tissue  of  the  mucosa  with  the  inter- 
cellular substance  of  the  epithelium.  The  stratified  columnar  epithelium 
is  formed  of  several  layers,  of  which  the  most  superficial  layer  is  ciliated  and 


FIG.  220. — Ciliary  Epithelium  of  the  Human  Trachea,  a,  Layer  of  longitudinally 
arranged  elastic  fibers;  &,  basement  membrane;  c,  deepest  cells,  circular  in  form;  d,  inter- 
mediate elongated  cells;  e,  outermost  layer  of  cells  fully  developed  and  bearing  cilia.  X 
350.  (Kolliker.) 

the  cells  often  branched  downward.  Many  of  the  superficial  cells  are  of  the 
goblet  variety.  In  the  deeper  part  of  the  mucosa  are  many  elastic  fibers 
between  which  lie  connective-tissue  corpuscles  and  capillary  blood  vessels. 

Numerous  mucous  glands  are  situated  on  the  exterior  and  in  the  substance 
of  the  fibrous  framework  of  the  trachea,  their  ducts  perforating  the  various 


FIG.  221. — Transverse  Section  of  a  Bronchus,  about  $  inch  in  Diameter,  e,  Epithelium 
(ciliated);  immediately  beneath  it  is  the  mucous  membrane  or  internal  fibrous  layer,  of 
varying  thickness;  m,  muscular  layer;  s,  m,  submucous  tissue;/,  fibrous  tissue;  c,  cartilage 
enclosed  within  the  layers  of  fibrous  tissue;  g,  mucous  gland.  (F.  E.  Schulze.) 

structures  which  form  the  wall  of  the  trachea,  and  opening  through  the 
mucous  membrane  into  the  cavity  of  the  trachea. 

The  two  bronchi  into  which  the  trachea  divides  resemble  the  trachea 
in  structure,  with  the  difference  that  in  them  there  is  a  distinct  layer  of  un- 
striped  muscle  arranged  circularly  beneath  the  mucous  membrane,  forming 


282  RESPIRATION 

the  muscularis  mucosa.  On  entering  the  substance  of  the  lungs  the  carti- 
laginous rings,  although  they  still  form  only  larger  or  smaller  segments  of 
a  circle,  are  no  longer  confined  to  the  front  and  sides  of  the  tubes,  but  are 
distributed  impartially  to  all  parts  of  their  circumference. 

The  bronchi  divide  and  subdivide  in  the  substance  of  the  lungs  into 
smaller  and  smaller  branches,  which  penetrate  into  every  part  of  the  organ 
until  at  length  they  end  in  the  smaller  subdivisions  of  the  lungs  called 
lobules. 

All  the  larger  branches  have  walls  formed  of  tough  membrane,  contain- 
ing portions  of  cartilaginous  rings,  by  which  they  are  held  open,  and  un- 
striped  muscular  fibers,  as  well  as  longitudinal  bundles  of  elastic  tissue. 
They  are  lined  by  mucous  membrane,  the  surface  of  which,  like  that  of  the 
larynx  and  trachea,  is  covered  with  ciliated  epithelium;  but  the  several 
layers  become  less  and  less  distinct  until  the  lining  consists  of  a  single  layer 
of  more  or  less  cubical  cells  covered  with  cilia,  figure  221.  The  mucous 
membrane  is  abundantly  provided  with  mucous  glands. 

As  the  bronchi  become  smaller  and  smaller  and  their  walls  thinner,  the 
cartilaginous  rings  become  fewer  and  more  irregular,  until  in  the  smaller 
bronchial  tubes  they  are  represented  only  by  minute  and  scattered  cartilag- 
inous flakes.  And  when  the  bronchi  by  successive  branches  are  reduced 
to  about  -fa  of  an  inch,  0.6  mm.,  in  diameter,  they  lose  their  cartilaginous  ele- 
ment altogether  and  their  walls  are  formed  only  of  a  tough,  fibrous,  elastic 
membrane  with  circular  muscular  fibers.  They  are  still  lined,  however, 
by  a  thin  mucous  membrane  with  ciliated  epithelium,  the  length  of  the 
cells  bearing  the  cilia  having  become  so  far  diminished  that  the  cells  are 
almost  cubical.  In  the  smaller  bronchi  the  circular  muscular  fibers  are 
relatively  more  abundant  than  in  the  larger  bronchi  and  form  a  distinct 
circular  coat. 

The  Lungs  and  Pleurae. — The  lungs  occupy  the  greater  portion  of 
the  thorax.  They  are  of  a  spongy  elastic  texture,  and  on  section  appear 
to  the  naked  eye  as  if  they  were  in  great  part  solid  organs,  except  where 
branches  of  the  open  bronchi  or  air-tubes  may  have  been  cut  across  and  show 
on  the  surface  of  the  section.  In  fact,  however,  the  lungs  are  hollow  organs 
composed  of  a  mass  of  air  cavities  all  of  which  communicate  finally  with 
the  common  air-tube,  the  trachea. 

Each  lung  is  enveloped  by  a  serous  membrane,  the  pleura,  which  ad- 
heres closely  to  its  surface  and  provides  it  with  its  smooth  and  slippery 
covering.  This  same  membrane  lines  the  inner  surface  of  the  chest  wall. 
The  continuity  of  this  membrane,  which  forms  a  closed  sac  as  in  the  case 
of  other  serous  membranes,  will  be  best  understood  by  reference  to  figure  222. 
The  appearance  of  a  space,  however,  between  the  pleura  which  covers  the 
lung,  visceral  layer,  and  that  which  lines  the  inner  surface  of  the  chest,  parietal 
layer,  is  inserted  in  the  drawing  only  for  the  sake  of  distinctness.  These 


THE    FINER    STRUCTURE    OF    THE    LUNG  283 

layers  are,  in  health,  everywhere  in  contact,  one  with  the  other;  and  between 
them  is  only  just  as  much  fluid  as  will  insure  frictionless  movement  in  their 
expansion  and  contraction. 

When  considering  the  subject  of  normal  respiration,  one  may  discard 
altogether  the  notion  of  the  existence  of  any  space  or  cavity  between  the 
lungs  and  the  wall  of  the  chest.  If,  however,  an  opening  be  made  so  as  to 
permit  air  or  fluid  to  enter  the  pleural  sac,  the  lung  in  virtue  of  its  elasticity 
recoils,  and  a  considerable  space  is  left  between  it  and  the  chest  wall.  In 
other  words,  the  natural  elasticity  of  the  lungs  would  cause  them  at  all  times 
to  contract  away  from  the  ribs  were  it  not  that  the  contraction  is  resisted  by 
atmospheric  pressure  which  bears  only  on  the  inner  surface  of  the  air-tubes 
and  air-cells. 

The  pulmonary  pleura  consists  of  an  outer  or  denser  layer  and  an  inner 
looser  tissue  in  which  there  is  a  lymph-canalicular  system.  Numerous 


FIG.  222. — Transverse  Section  of  the  Chest. 

lymphatics  are  to  be  met  with,  which  form  a  dense  plexus  of  vessels,  many 
of  which  contain  valves.  They  are  simple  endothelial  tubes,  and  take  origin 
in  the  lymph-canalicular  system  of  the  pleura  proper.  Scattered  bundles 
of  unstriped  muscular  fiber  occur  in  the  pulmonary  pleura.  They  are  es- 
pecially strongly  developed  on  the  anterior  and  internal  surfaces  of  the  lungs, 
the  parts  which  move  most  freely  in  respiration.  Their  function  is  doubt- 
less to  aid  in  expiration. 

The  Finer  Structure  of  the  Lung. — Each  lung  is  partially  subdi- 
vided into  separate  portions  called  lobes:  the  right  lung  into  three  lobes 
and  the  left  into  two.  Each  of  these  lobes,  again,  is  composed  of  a  large  num- 
ber of  minute  parts,  called  lobules.  Each  pulmonary  lobule  may  be  con- 
sidered to  be  a  lung  in  miniature,  consisting,  as  it  does,  of  a  branch  of  the 
bronchial  tube,  of  air-cells,  blood  vessels,  nerves,  and  lymphatics,  with  a 
small  amount  of  areolar  tissue. 


284  RESPIRATION 

On  entering  a  lobule,  the  small  bronchial  tube,  the  structure  of  which 
has  just  been  described,  a,  figure  223,  divides  and  subdivides;  its  walls  at 
the  same  time  becoming  thinner  and  thinner,  until  at  length  they  are  formed 
only  of  a  thin  membrane  of  areolar  and  elastic  tissue,  lined  by  a  layer  of 
squamous  epithelium,  no  longer  provided  with  cilia.  At  the  same  time  they 
are  altered  in  shape;  each  of  the  minute  terminal  branches  widening  out 
funnel-wise,  and  its  walls  being  pouched  out  irregularly  into  small  saccular 
dilatations,  called  air-cells,  figure  223,  b.  Such  a  funnel-shaped  terminal 
branch  of  the  bronchial  tube,  with  its  group  of  pouches  or  air-cells,  has  been 
called  an  infundibulum,  figures  223  and  224,  and  the  irregular  oblong  space 
in  its  center,  with  which  the  air-cells  communicate,  an  intercellular  passage. 


FIG.  223.  FIG.  224. 

FIG.  223. — Terminal  Branch  of  a  Bronchial  Tube,  with  its  Infundibula  and  Air-cells, 
from  the  Margin  of  the  Lung  Injected  with  Quicksilver;  Monkey,  a,  Terminal  bronchial 
twig;  6,  6,  infundibula  and  air-cells.  X  10.  (F.  E.  Schulze.) 

FIG.  224. — Two  Small  Infundibula,  a,  a,  with  air-cells,  b,  b,  and  the  ultimate  bronchial 
tubes,  c,  c,  with  which  the  air-cells  communicate.  From  a  new-born  child.  (Kolliker.) 


An  inflated  and  dried  turtle's  lung  illustrates  the  homologue  of  a  lobule. 
Such  a  preparation  can  be  cut  across  to  illustrate  the  intercellular  passage, 
the  infundibulum,  and  the  air-cells. 

The  air-cells,  or  air-vessels,  are  sometimes  placed  singly,  like  recesses 
from  the  intercellular  passage,  but  more  often  they  are  arranged  in  groups 
or  even  rows,  like  minute  sacculated  tubes,  so  that  a  short  series  of  vesicles 
all  communicating  with  one  another  open  by  a  common  orifice  into  the  tube. 
The  vesicles  are  of  various  forms  according  to  the  mutual  pressure  to  which 
they  are  subject.  Their  walls  are  nearly  in  contact,  and  they  vary  from  o .  3 
to  0.5  mm.  in  diameter.  Their  walls  are  formed  of  fine  membrane  similar 
to  that  of  the  intercellular  passages  and  continuous  with  it.  The  membrane 
is  folded  on  itself  so  as  to  form  a  sharp-edged  border  at  each  circular  orifice 


THE  FINER  STRUCTURE  OF  THE  LUNG 


28; 


of  communication  between  contiguous  air-vesicles,  or  between  the  vesicles 
and  the  bronchial  passages.  Numerous  fibers  of  elastic  tissue  are  spread 
out  in  the  walls  between  contiguous  air-cells,  and  many  of  these  are  attached 
to  the  outer  surface  of  the  wall  of  which  each  cell  is  composed,  imparting  to 
it  additional  strength  and  the  power  of  recoil  after  distention. 

The  air-cells  are  lined  by  a  layer  of  epithelium,  figure  225,  the  cells  of 
which  are  very  thin  and  plate-like.  The  thin  epithelial  membrane  is  free  on 
one  side,  where  it  comes  in  contact  with  the  air  of  the  lungs,  but  on  the  other 


FIG.  225. — From  a  Section  of  the  Lung  of  a  Cat,  Stained  with  Silver  Nitrate.  A.  D, 
Alveolar  duct  or  intercellular  passage;  S,  alveolar  septa,  N,  alveoli  or  air-cells,  lined  with 
large,  flat,  nucleated  cells,  with  some  smaller  polyhedral  nucleated  cells;  M,  unstriped 
muscular  fibers.  Circular  muscular  fibers  are  seen  surrounding  the  interior  of  the  alveolar 
duct,  and  at  one  part  is  seen  a  group  of  small  polyhedral  cells  continued  from  the  bronchus. 
(Klein  and  Noble  Smith.) 


side  a  network  of  pulmonary  capillaries  is  spread  out  so  densely,  figure  226 
that  the  interspaces  or  meshes  are  even  narrower  than  the  vessels.  These 
are  on  an  average  3^Vo  °f  an  inch,  or  8  micromillimeters,  in  diameter.  Be- 
tween the  atmospheric  air-cells  and  the  blood  in  these  vessels,  nothing  in- 
tervenes but  the  thin  walls  of  the  cells  and  capillaries.  The  exposure  of  the 
blood  to  the  air  is  the  more  complete  because  the  wall  between  contiguous 
air-cells,  and  often  the  spaces  between  the  walls  of  the  same,  contain  only 
a  single  layer  of  capillaries  both  sides  of  which  are  at  once  exposed  to  the  air. 

The  air-vesicles  situated  nearest  to  the  center  of  the  lung  are  smaller 
and  their  networks  of  capillaries  are  closer  than  those  nearer  to  the  circum- 


'286  RESPIRATION 

ference.  The  vesicles  of  adjacent  lobules  do  not  communicate.  Those  of 
the  same  lobule  or  proceeding  from  the  same  intercellular  passage  com- 
municate, as  a  general  rule,  only  near  angles  of  bifurcation,  so  that  when  any 
bronchial  tube  is  closed  or  obstructed  the  supply  of  air  is  lost  for  all  the  blood 
vessels  of  that  lobule  and  its  branches. 

Blood  Supply. — The  lungs  receive  blood  from  two  sources:  a,  the 
pulmonary  artery;  b,  the  bronchial  arteries.  The  former  conveys  venous 
blood  to  the  lungs  for  its  oxidation,  and  this  blood  takes  no  share  in  the 


FIG.  226. — Section  of  Injected  Lung,  Including  Several  Contiguous  Alveoli.  (F.  E. 
Schulze.)  Highly  magnified,  a,  a,  Free  edges  of  alveoli;  c,  c,  partitions  between  neighbor- 
ing alveoli,  seen  in  section;  &,  small  arterial  branch  giving  off  capillaries  to  the  alveoli. 
The  looping  of  the  vessels  to  either  side  of  the  partitions  is  well  exhibited.  Between  the 
capillaries  is  seen  the  homogeneous  alveolar  wall  with  nuclei  of  connective-tissue  corpuscles 
and  elastic  fibers. 


nutrition  of  the  deeper  pulmonary  tissues  through  which  it  passes.  The 
branches  of  the  bronchial  arteries  are  nutrient  arteries  which  ramify  in  the 
walls  of  the  bronchi,  in  the  walls  of  the  larger  pulmonary  vessels,  and  in  the 
interlobular  connective  tissue,  etc.  The  blood  of  the  bronchial  vessels  is  re- 
turned chiefly  through  the  bronchial,  but  partly  through  the  pulmonary,  veins. 
Lymphatics. — The  lymphatics  are  arranged  in  three  sets:  i.  Ir- 
regular lacunae  in  the  walls  of  the  alveoli  or  air-cells.  The  lymphatic  vessels 
which  lead  from  these  accompany  the  pulmonary  vessels  toward  the  root 
of  the  lung.  2,  Irregular  anastomosing  spaces  in  the  walls  of  the  bronchi. 
3,  Lymph  spaces  in  the  pulmonary  pleura.  The  lymphatic  vessels  from  all 


THE   MOVEMENTS    OF   THE   RESPIRATORY   MECHANISM  287 

these  irregular  sinuses  pass  in  toward  the  root  of  the  lung  to  reach  the  bron- 
chial glands. 

Nerves. — The  nerves  of  the  lung  are  to  be  traced  from  the  anterior 
and  posterior  pulmonary  plexuses,  which  are  formed  by  branches  of  the 
vagus  and  sympathetic.  The  nerves  follow  the  course  of  the  blood  vessels 
and  bronchi,  and  many  small  ganglia  are  situated  in  the  walls  of  the  latter. 


FIG.  227. — Capillary  Network  of  the  Pulmonary  Blood  Vessels  in  the  Human  Lung.     X  60. 

(Kolliker.) 

THE  MOVEMENTS  OF  THE  RESPIRATORY  MECHANISM. 

Respiratory  movement  consists  of  the  alternate  expansion  and  contrac- 
tion of  the  thorax,  by  means  of  which  air  is  drawn  into,  or  expelled  from, 
the  lungs. 

A  movement  of  the  side  walls  or  floor  of  the  chest  to  increase  its  diameter 
or  length  will  enlarge  the  capacity  of  the  interior.  By  such  an  increase  of 
capacity  there  will  be  of  course  a  diminution  of  the  pressure  of  the  air  in  the 
lungs,  and  a  fresh  quantity  of  air  will  enter  through  the  larynx  and  trachea 
to  equalize  the  pressure  on  the  inside  and  outside  of  the  chest.  This  move- 
ment is  called  inspiration. 

The  movement  which  diminishes  the  capacity  of  the  chest  and  increases 
the  pressure  in  the  interior  expels  air  until  the  pressure  within  and  that  with- 
out the  chest  are  again  equal.  This  movement  is  called  expiration.  In  both 
cases  the  air  passes  through  the  trachea  and  larynx,  whether  in  entering  or 
leaving  the  lungs,  there  being  no  other  communication  with  the  exterior  of 
the  body.  And  the  lung,  for  the  same  reason,  remains  closely  in  contact 
with  the  walls  and  floor  of  the  chest  under  all  the  circumstances  described. 
To  speak  of  expansion  of  the  chest  is  to  speak  also  of  expansion  of  the  lung, 
and  vice  versa. 


288 


RESPIRATION 


Inspiration. — The  enlargement  of  the  chest  during  inspiration  is  due  to 
muscular  action,  which  brings  about  an  increase  in  the  size  of  the  chest  cavity 
through  the  contraction  of  the  inspiratory  muscles,  the  role  played  by  the 
lungs  being  a  passive  one.  The  chest  cavity  is  increased  in  its  three  axes, 
the  vertical,  lateral,  and  antero-posterior  diameters.  The  muscles  engaged 
in  ordinary  inspiration  are:  the  diaphragma,  the  intercostales  externi,  and 
the  scaleni  and  levatores  costarum.  During  forced  inspiration  every 
muscle  is  brought  into  play  which  by  its  contraction  tends  to  elevate  the  ribs 

and  sternum  or  which  will  fix  points  against 
which  these  muscles  can  act.  This  includes 
almost  every  muscle  of  the  trunk  and  neck. 
Changes  in  the  vertical  diameter  are  due, 
first,  to  the  contraction  of  the  diaphragm. 
This  muscle  has  the  shape  of  a  flattened 
dome,  its  highest  point  being  the  central 
tendon.  While  passive  its  lower  portions 
are  in  apposition  with  the  chest  walls,  figure 
228,  7.  On  contraction,  the  dome  is  pulled 
downward  and  the  lower  portion  is  pulled 
away  from  the  chest  walls,  the  downward 
displacement  varying  from  6  to  12  mm.  in 
normal  respiration,  and  in  forced  respira- 
tion may  amount  to  as  much  as  45  mm. 
The  tendency  of  the  diaphragm  to  pull  the 
lower  ribs  and  lower  part  of  the  sternum 

inward  is  counteracted  by  the  outward  pressure  of  the  abdominal  viscera, 
and  by  the  action  of  the  quadrati  lumbori,  which  by  their  attachment  to 
the  last  ribs  fix  these  and,  in  case  of  deep  inspiration,  may  even  pull  them 
downward.  The  serrati  postici  inferiores  also  aid,  being  attached  to  the 
four  lower  ribs. 

Changes  in  the  lateral  and  antero-posterior  diameters  are  effected  by  the 
raising  of  the  ribs,  which  are  attached  very  obliquely  to  the  spine  and  sternum. 
The  elevation  of  the  ribs  takes  place  both  in  front  and  at  the  sides — the 
hinder  ends  being  prevented  from  performing  any  upward  movement  by 
their  pivot  attachment  to  the  spine.  The  movement  of  the  front  extremities 
of  the  ribs  is  of  necessity  limited  by  an  upward  and  forward  movement  of  the 
sternum  to  which  they  are  attached,  the  movement  being  greater  at  the  lower 
end  than  at  the  upper  end  of  the  sternum. 

The  axes  of  rotation  in  these  movements  are  two:  one  corresponding 
with  a  line  drawn  through  the  two  articulations  which  the  rib  forms  with 
the  spine,  o,  b,  figure  230,  and  the  other  with  a  line  drawn  from  one  of  these 
(head  of  rib)  to  the  sternum,  A  B,  figure  230;  the  motion  of  the  rib  around 
the  latter  axis  being  somewhat  after  the  fashion  of  raising  the  handle  of  a 


FIG.  228. — Schematic  Repre- 
sentation of  Diaphragm.  In  ex- 
piration (7),  quiet  inspiration 
(77),  and  deep  inspiration  (///). 
(After  Schaffer.) 


INSPIRATION 


289 


bucket.  The  elevation  of  the  ribs  is  accompanied  by  a  slight  opening  out  of 
the  angle  which  the  bony  part  forms  with  its  cartilage,  and  thus  an  additional 
means  is  provided  for  increasing  the  antero-posterior  diameter  of  the  chest. 
The  movements  of  all  the  ribs  except  the  twelfth  consist  of  a  rotation  up- 


CEsophagus 
L*fl  subcUvten  artery 
Left  common  carotid  artery 

Left  superior  intercostal  vein 
Left  innominate  vei 


Parietal 
pleura 
(cut  edge) 


ihagus 


Diaphrasm 


FIG.  229. — Thorax  from  the  Left,  Showing  Left  Pleural  Sac,  and  the  Diaphragm.     The 
lung  is  removed.     (Cunningham.) 


ward,  forward,  and  outward.     The  twelfth  presents  only  rotation  down- 
ward and  backward. 

The  muscles  involved  in  these  movements  of  the  ribs  are  the  external 
intercostals  and  the  part  of  the  internal  intercostals  situated  between  the 
costal  cartilages.  Their  action  is  to  widen  the  intercostal  spaces.  The 
scaleni  fix  the  first  and  second  ribs,  thereby  making  a  fixed  point  of  action 


RESPIRATION 


for  the  other  muscles  involved.  The  serrati  postici  superiores  assist  the  above 
and  also  raise  the  third,  fourth,  and  fifth  ribs.  The  levatores  costarum  longi 
and  brevi  elevate  and  evert  all  the  ribs  from  the  first  to  the  tenth. 

In  extraordinary  or  forced  inspiration,  which  may  be  due  either  to  violent 
exercise  or  to  interference  with  the  due  entrance  of  air  into  the  lungs,  all  the 
above  muscles  act  more  strongly.  The  diaphragm  descends  lower,  the 
scaleni  raise  the  first  and  second  ribs  instead  of  merely  fixing  them,  as  in 
ordinary  respiration,  as  do  also  the  sterno-cleido-mastoids.  These,  together 
with  the  sacro-spinales  which  straighten  the  spine,  increase  the  vertical 
diameter.  The  trapezii  and  the  rhomboidii  assist  in  increasing  the  antero- 


FiG.  230. — Diagram  of  Axes  of  Movement  of  Ribs. 

posterior  and  lateral  diameters  by  fixing  the  shoulders  and  thus  giving  a 
fixed  point  for  the  action  of  the  pectorales  and  latissimi  dorsi. 

The  enlargement  of  the  chest  during  inspiration  presents  peculiarities 
in  different  persons.  In  children  of  both  sexes  the  principal  muscle  in- 
volved seems  to  be  the  diaphragm,  and  this  type  of  breathing  is  known  as 
abdominal  breathing.  In  men,  the  chest  and  sternum,  together  with  the 
front  wall  of  the  abdomen,  are  subject  to  a  wide  movement;  this  type  of 
breathing  is  called  the  inferior  costal.  In  women,  the  movement  appears 
less  extensive  in  the  lower  and  more  extensive  in  the  upper  part  of  the  chest, 
which  is  called  the  superior  costal  type.  This  has  been  shown  to  be  due 
rather  to  mode  of  dress  than  to  a  real  difference  in  the  sexes  (Mosher). 

Expiration. — Quiet  expiration  is  a  passive  act  due  to  the  return  of 
the  thorax  and  its  contained  lungs  to  their  normal  position  when  the  mus- 
cles involved  in  inspiration  relax.  This  elastic  recoil  is  sufficient  in  ordinary 
quiet  breathing  to  expel  air  from  the  lungs.  In  forced  expiration,  however, 


RECORDING    RESPIRATORY    MOVEMENTS  2QI 

which  may  occur  to  a  slight  degree  in  speaking,  singing,  etc.,  as  well  as  in 
the  case  of  many  involuntary  and  reflex  acts,  such  as  coughing,  sneezing, 
etc.,  other  muscles  are  involved.  Of  these  the  principal  are  the  abdominal 
muscles,  obliquus  externus  and  internus,  rectus  abdominis,  transversus  ab- 
dominis  and  pyramidalis .  These  act,  first,  by  pressing  the  abdominal 
viscera  against  the  diaphragm  and  thereby  forcing  it  up,  their  descent  into 
the  pelvic  cavity  being  prevented;  second,  by  their  attachments  to  the  lower 
ribs  and  cartilages,  the  muscles  draw  these  downward  and  inward,  thereby 
lessening  the  size  of  the  thoracic  cavity;  lastly,  by  their  contraction,  they 
form  a  fixed  point  for  the  action  of  that  part  of  the  internal  intercostals, 
not  involved  in  inspiration,  to  approximate  the  ribs. 

When  by  the  efforts  of  the  expiratory  muscles  the  chest  has  been  squeezed 
to  less  than  its  average  diameters,  it  again,  on  relaxation  of  the  muscles, 
returns  to  the  normal  dimensions  by  virtue  of  its  elasticity.  The  construc- 
tion of  the  chest  walls,  therefore,  admirably  adapts  them  for  recoiling  against 
and  resisting  as  well  undue  contraction  as  undue  dilatation. 

Respiratory  Movements  of  the  Nostrils  and  of  the  Glottis. — During 
the  action  of  the  inspiratory  muscles  which  directly  draw  air  into  the  chest, 
those  which  guard  the  opening  through  which  the  air  enters  are  also  active. 
In  hurried  breathing  the  dilatation  of  the  nostrils  is  well  seen,  although 
under  ordinary  conditions  it  may  not  be  noticeable.  The  opening  at  the 
upper  part  of  the  larynx,  however,  the  rima  glottidis,  is  dilated  at  each  in- 
spiration for  the  more  ready  passage  of  air,  and  becomes  smaller  at  each 
expiration;  its  condition,  therefore,  corresponds  during  respiration  with 
that  of  the  walls  of  the  chest.  There  is  a  further  likeness  between  the  two 
acts  in  that,  under  ordinary  circumstances,  the  dilatation  of  the  rima  glot- 
tidis is  a  muscular  act  and  its  contraction  chiefly  an  elastic  recoil;  although, 
under  various  special  conditions  to  be  hereafter  mentioned,  there  may  be 
considerable  muscular  contraction  exercised. 

Methods  of  Recording  Respiratory  Movements. — The  movements  of  respira- 
tion may  be  recorded  graphically  in  several  ways.  The  ordinary  method  is  to 
introduce  a  tube  into  the  trachea  of  an  animal,  and  to  connect  this  tube  by 
some  gutta-percha  tubing  with  a  T-piece,  the  side  branch  of  which  is  connected 
with  a  Marey's  tambour,  which  may  be  made  to  write  on  a  recording  surface, 
figure  156.  If  the  tube  attached  to  the  free  limb  of  the  T-piece  be  partially 
closed  with  a  screw  compress,  the  movements  of  inspiration  and  expiration  are 
larger  than  if  it  were  open.  The  alteration  of  the  pressure  within  the  lungs  on 
inspiration  and  expiration  is  shown  by  the  movement  of  the  column  of  air  in 
the  trachea  and  in  its  extension  to  the  T-piece.  By  these  means  a  record  of  the 
respiratory  movements  may  be  obtained  in  experimental  animals. 

Various  instruments  have  been  devised  for  recording  the  movements  of 
the  chest  by  application  of  apparatus  to  the  exterior.  Such  is  the  stethometer 
of  Burdon-Sanderson,  figure  233.  This  consists  of  a  frame  formed  of  two 
parallel  steel  bars  joined  by  a  third  at  one  end.  At  the  free  end  of  the  bars 
is  attached  a  leather  strap,  by  means  of  which  the  apparatus  may  be  suspended 


2Q2 


RESPIRATION 


from  the  neck.  Attached  to  the  inner  end  of  one  bar  is  a  tambour  and  ivory 
button,  to  the  end  of  the  other  an  ivory  button.  The  apparatus  is  suspended 
with  the  transverse  bar. posteriorly,  the  button  of  the  tambour  is  placed  on  the 
part  of  the  chest  the  movement  of  which  it  is  desired  to  record,  and  the  other  but- 
ton is  made  to  press  upon  the  corresponding  side  of  the  chest,  so  that  the  chest 


FIG.  231. — Stethograph  or  Pneumograph.  h,  Tambour  fixed  at  right  angles  to  plate 
of  steel,  /;  c  and  d,  arms  by  which  instrument  is  attached  to  chest  by  belt,  e.  When  the 
chest  expands,  the  arms  are  pulled  asunder,  which  bends  the  steel  plate,  and  the  tambour  is 
affected  by  the  pressure  of  b,  which  is  attached  to  it  on  the  one  hand,  and  to  the  upright  in 
connection  with  horizontal  screw,  g.  (Modified  from  Marey's  instrument.) 

is  held  as  between  a  pair  of  calipers.  The  receiving  tambour  is  connected 
through  a  T-piece  with  a  recording  tambour  of  Marey's  and  with  a  bulb  by 
means  of  which  air  can  be  squeezed  into  the  cavity  of  the  typanum.  When 
adjusted  the  tube  connected  with  the  air  ball  is  shut  off  by  means  of  a  screw 
clamp.  The  movement  of  the  chest  is  thus  communicated  to  the  recording 
tambour. 


FIG.  232. — Tracing  of  Thoracic  Respiratory  Movements  obtained  by  means  of 
Marey's  Pneumograph.  A  whole  respiratory  phase  is  comprised  between  a  and  a; 
inspiration  during  which  the  lever  descends,  extending  from  a  to  b,  and  expiration  from  b 
to  a.  The  undulations  at  c  are  caused  by  the  heart's  beat.  (Foster.) 

A  simpler  form  of  this  apparatus,  called  a  pneumograph  or  stethograph, 
consists  of  a  thick  india-rubber  bag  of  elliptical  shape  about  three  inches  long, 
to  one  end  of  which  a  rigid  gutta-percha  tube  is  attached.  This  bag  may  be 
fixed  at  any  required  place  on  the  chest  by  means  of  a  strap  and  buckle.  By 
means  of  the  gutta-percha  tube  the  variations  of  the  pressure  of  air  in  the  bag, 


RELATIVE    TIME    OF    INSPIRATION    AND    EXPIRATION 


293 


produced  by  the  movements  of  the  chest,  are  communicated  to  a  recording 
tambour.  This  principle  is  applied  in  a  modified  form  in  Marey's  pneumo- 
graph,  figure  231. 

The  variations  of  intrapleural  pressure  may  be  recorded  by  introducing  a 
cannula  into  the  pleural  or  pericardial  cavity.  The  cannula  should  be  pre- 
viously connected  with  a  mercury  or  other  form  of  manometer  by  tubing 
filled  with  physiological  saline. 


Tambour. 
Ivory  button. 


Tube  to  commu- 
nicate with  re- 
cording tam- 
bour. 


Ball  to  fill  appa-  _. 
ratua  with  air. 

FIG.  233. — Stethometer.     (Burdon-Sanderson.) 

Finally,  it  has  been  found  possible  in  various  ways  to  record  the  diaphrag- 
matic movements.  This  can  be  done  by  inserting  a  receiving  tambour  into 
the  abdomen  below  the  diaphragm,  by  the  insertion  of  needles  into  different 
parts  of  the  diaphragm  and  recording  the  movement  of  the  free  ends  of  needles 
about  the  fulcrum  formed  where  the  chest  wall  is  pierced,  or  by  recording  the 
contraction  of  isolated  strips  of  the  diaphragm  directly.  These  records  all 
give  an  accurate  picture  of  the  movements  of  the  diaphragm. 

The  Relative  Time  of  Inspiration  and  Expiration  and  the  Respira- 
tory Movement. — The  acts  of  inspiration  and  expiration  take  up,  under 
ordinary  circumstances,  a  nearly  equal  time.  The  time  of  inspiration, 
however,  especially  in  women  and  children,  is  a  little  shorter  than  that  of 
expiration,  and  there  is  commonly  a  very  slight  pause  between  the  end  of 
expiration  and  the  beginning  of  the  next  inspiration,  see  figure  232.  The 
ratio  of  the  respiratory  rhythm  may  be  thus  expressed: 

Inspiration 6 

Expiration 7  to  8 

Pause Very  slight 


2Q4 


RESPIRATION 


If  the  ear  be  placed  in  contact  with  the  wall  of  the  chest  or  be  separated 
from  it  only  by  a  good  conductor  of  sound  or  a  stethoscope,  a  faint  respiratory 
murmur  is  heard  during  inspiration.  This  sound  varies  somewhat  in  dif- 
ferent parts,  being  loudest  or  coarsest  in  the  neighborhood  of  the  trachea  and 
large  bronchi  (tracheal  and  bronchial  breathing),  and  fading  off  into  a  faint 
sighing  as  the  ear  is  placed  at  a  distance  from  these  (vesicular  breathing). 
It  is  heard  best  in  children.  In  them  a  faint  murmur  is  heard  in  expiration 
also.  The  cause  of  the  vesicular  murmur  has  received  various  explanations. 
Most  observers  hold  that  the  sound  is  produced  in  the  glottis  and  larger 


FIG.  234. — Tracing  of  the  Normal  Diaphragm  Respirations  of  the  Rabbit,  a,  With 
quick  movement  of  drum;  b,  with  slow  movement;  /,  inspiration;  E,  expiration.  To  be 
read  from  left  to  right.  (Marckwald.) 


bronchial  tubes,  but  that  it  is  modified  in  its  passage  to  the  pulmonary 
alveoli.  In  disease  of  the  lungs  the  vesicular  murmur  undergoes  various 
modifications,  for  a  description  of  which  One  must  consult  text-books  on 
physical  diagnosis. 

The  Quantity  of  Air  Breathed. — Tidal  air  is  the  quantity  of  air 
which  is  habitually  and  almost  uniformly  changed  in  each  act  of  breathing. 
In  a  healthy  adult  man  it  is  about  30  cubic  inches,  or  about  500  cc.  or  half 
a  liter.  In  college  students  the  tidal  air  is  somewhat  less,  varying  from  300 
to  400  cc.  while  at  rest. 

The  complemental  air  is  the  quantity  of  air  which  can  be  drawn  into  the 
lungs  by  the  deepest  inspiration  over  and  above  that  which  is  in  the  lungs 
at  the  end  of  an  ordinary  inspiration.  Its  amount  varies,  but  may  be  reck- 
oned as  100  cubic  inches,  or  about  1,600  cc. 


QUANTITY    OF    AIR   BREATHED 


295 


The  reserve  air  is  that  which  may  be  expelled  by  a  forcible  and  deeper 
expiration,  after  an  ordinary  expiration,  such  as  that  which  expels  the 
tidal  air.  The  reserve  air  amounts  to  from  1,200  to  1,500  cc.  This  is  also 
termed  the  supplemental  air. 


FIG.  235. — Photograph  of  the  Sanborn  Company  form  of  spirometer.  The  grad- 
uated disc  records  the  volume  of  air  exhaled.  In  using  the  spirometer  the  reservoir  is 
first  filled  with  water  to  form  a  water  seal  for  the  air  chamber.  To  perform  a  respi- 
ratory volume  test  the  instrument  is  set  automatically  at  zero,  though  it  is  well  to 
begin  with  the  air  chamber  empty.  Place  the  previously  sterilized  mouth  piece  between 
the  lips  and  if  necessary,  close  the  nostrils  with  pinch  cock  or  with  the  hand.  The 
receiving  chamber  is  delicately  balanced  and  as  the  breath  is  forced  through  the  tube  into 
this  bell  it  rises,  recording  the  movement  on  the  circular  index,  from  the  scale  of  which 
the  volume  is  read  off  directly.  For  very  accurate  comparative  determinations  the 
expired  air  should  be  allowed  to  stand  long  enough  to  come  to  constant  temperature, 
and  corrections  for  variations  from  the  standard  temperature  and  pressure  should  be 
made.  This  is  not  necessary  in  the  routine  laboratory  and  gymnasium  measurements. 

The  residual  air  is  the  quantity  which  still  remains  in  the  lungs  after 
the  most  violent  expiratory  effort.  Its  amount  depends  in  great  measure 
on  the  absolute  size  of  the  chest,  but  may  be  estimated  at  about  i  ,000  cc. 

tO   1,200  CC. 

The  quantity  of  air  breathed  per  minute,  called  the  minute  volume, 
varies  in  the  adult  at  rest  according  to  size.  But  the  average  may  be  set 
down  as  between  5  and  8  liters.  In  24  hours  this  would  amount  to  from 


296  RESPIRATION 

7,200  to  11,500  liters.  However,  many  factors  lead  to  great  variations  in 
this  volume.  Vigorous  exercise  will  increase  the  minute  volume  to  12  to 
15  or  more  liters  per  minute.  Breathing  a  rarefied  air  will  produce  the 
same  result,  as  in  mountain  climbing  or  aviation.  The  lack  of  oxygen  or 
anoxemia  augments  the  minute  volume  both  by  accelerating  the  respira- 
tory rate  and  increasing  the  depth.  Excess  of  carbon  dioxide,  or 
after  moderate  carbon  monoxide  poisoning,  similar  response  is  given, 
though  the  rate  is  more  vigorously  affected  on  breathing  air  rich  in  carbon 
dioxide. 

The  Respiratory  Capacity. — The  greatest  respiratory  capacity  or  vital 
capacity  of  the  chest  is  indicated  by  the  quantity  of  air  which  a  person  can 
expel  from  his  lungs  by  a  forcible  expiration  after  the  deepest  possible  in- 
spiration. The  vital  capacity  is  the  sum  of  the  reserve,  tidal,  and  comple- 
mental  airs.  It  expresses  the  power  which  a  person  has  of  breathing  in  the 
emergencies  of  active  exercise,  violence,  and  disease.  The  average 
capacity  of  an  adult,  at  15.4°  C.  (60°  F.),  is  about  225  to  250  cc.,  or 
3,500  to  4,000  cc.  In  healthy  men,  the  respiratory  capacity  varies  chiefly 
with  the  stature,  weight,  and  age. 

Circumstances  Affecting  the  Amount  of  Respiratory  Capacity. — John 
Hutchinson  states  that  for  every  centimeter  of  height  above  the  standard 
the  respiratory  capacity  is  increased,  on  an  average,  by  50  cc. 

The  influence  of  weight  on  the  capacity  of  respiration  is  less  manifest, 
and  considerably  less  than  that  of  height.  It  is  difficult  to  arrive  at  any 
definite  conclusions  on  this  point,  because  the  natural  average  weight  of  a 
healthy  man  in  relation  to  stature  has  not  yet  been  determined. 

The  capacity  appears  to  be  increased  by  age  from  about  the  fifteenth 
to  the  thirty-fifth  year,  at  the  rate  of  80  cc.  per  year;  from  thirty- 
five  to  sixty-five  it  diminishes  at  the  rate  of  about  25  cc.  per  year;  so 
that  the  capacity  of  respiration  of  a  man  sixty  years  old  would  be  about 
480  cc.  less  than  that  of  a  man  forty  years  old,  of  the  same  height  and 
weight. 

The  number  of  respirations  in  a  healthy  adult  person  usually  ranges 
from  14  to  1 8  per  minute.  It  is  greater  in  infancy  and  childhood.  It 
varies  also  much  according  to  different  circumstances,  such  as  exercise  or 
rest,  health  or  disease,  etc.  Variations  in  the  number  of  respirations  corre- 
spond ordinarily  with  similar  variations  in  the  pulsations  of  the  heart.  In 
health  the  proportion  is  about  i  to  4,  or  i  to  5 ;  and  when  the  rapidity  of 
the  heart's  action  is  increased,  that  of  the  chest  movement  is  commonly 
increased  also,  but  not  in  every  case  in  equal  proportion.  It  happens 
occasionally  in  disease,  especially  of  the  lungs  or  air-passages,  that 
the  number  of  respiratory  acts  increases  in  quicker  proportion  than  the 
beats  of  the  pulse;  and,  in  other  affections,  much  more  commonly, 


COMPOSITION    OF    THE    ATMOSPHERE  297 

that  the  number  of  the  pulses  is  greater  in  proportion  than  that  of  the 
respirations. 

The  Force  of  Inspiratory  and  Expiratory  Muscles. — The  force  which 
the  inspiratory  muscles  are  capable  of  exerting  on  the  chest  is  greatest 
in  muscular  individuals  of  the  mean  height  of  about  five  feet  seven  or 
eight  inches  and  is  equal  to  a  column  of  two  and  a  half  to  three  inches  of 
mercury.  The  force  manifested  in  the  strongest  expiratory  acts  is,  on  the 
average,  one-third  greater  than  that  exercised  in  inspiration.  But  this 
difference  is  in  a  great  measure  due  to  the  power  exerted  by  the  elastic 
reaction  of  the  walls  of  the  chest;  and  it  is  also  much  influenced  by  the 
disproportionate  strength  which  the  expiratory  muscles  attain  from  their 
being  called  into  use  for  other  purposes  than  that  of  simple  expiration. 

Within  the  limits  of  ordinary  tranquil  respiration  the  elastic  resilience 
of  the  walls  of  the  chest  favors  inspiration.  It  is  only  in  deep  inspiration 
that  the  ribs  and  rib  cartilages  offer  an  opposing  force  to  dilatation.  In 
other  words,  the  elastic  resilience  of  the  lungs,  at  the  end  of  an  act  of 
ordinary  exhalation  has  drawn  the  chest  walls  within  the  limits  of  their 
normal  degree  of  expansion.  Under  all  circumstances,  of  course,  the 
elastic  tissue  of  the  lungs  opposes  inspiration  and  favors  expiration. 

It  is  possible  that  the  contractile  power  which  the  bronchial  tubes  and 
air- vesicles  possess,  by  means  of  their  muscular  fibers  may  assist  in  expiration. 
But  it  is  more  likely  that  its  chief  purpose  is  to  regulate  and  adapt,  in  some 
measure,  the  quantity  of  air  admitted  to  the  lungs,  and  to  each  part  of  them, 
according  to  the  supply  of  blood.  The  muscular  tissue  contracts  upon  and 
gradually  expels  collections  of  mucus,  which  may  have  accumulated  within 
the  tubes,  and  which  cannot  be  ejected  by  forced  expiratory  efforts,  owing 
to  collapse  or  other  morbid  condition  of  the  portion  of  lung  connected  with 
the  obstructed  tubes  (Gardner).  Apart  from  any  of  the  before- mentioned 
functions,  the  presence  of  muscular  fiber  in  the  walls  of  a  hollow  viscus,  such 
as  a  lung,  is  only  what  might  be  expected  from  analogy  with  other  organs. 
Subject  as  the  lungs  are  to  such  great  variation  in  size,  it  might  be  antici- 
pated that  the  elastic  tissue,  which  enters  so  largely  into  their  composition, 
would  be  supplemented  by  the  presence  of  much  muscular  fiber. 

RESPIRATORY  CHANGES  IN  THE  AIR  BREATHED. 


Composition  of  the  Atmosphere. — The  atmosphere  we  breathe  has, 
in  every  situation  in  which  it  has  been  examined  in  its  natural  state,  a  nearly 
uniform  composition.  It  is  a  mixture  of  oxygen,  nitrogen,  carbon  dioxide, 
and  watery  vapor,  with,  commonly,  traces  of  other  gases,  as  argon,  ammo- 
nia, sulphureted  hydrogen,  etc.  Of  every  100  volumes  of  pure  atmospheric 
air,  79  volumes,  on  an  average,  consist  of  nitrogen  and  argon,  the  remaining 


RESPIRATION 

21  of  oxygen.  The  proportion  of  carbon  dioxide  is  extremely  small;  10,000 
volumes  of  atmospheric  air  contain  only  about  4  of  that  gas. 

The  quantity  of  watery  vapor  varies  greatly  according  to  the  tempera- 
ture and  other  circumstances,  but  the  atmosphere  is  never  without  some. 
In  this  country  the  average  quantity  of  watery  vapor  in  the  atmosphere 
varies  greatly  according  to  the  region.  In  some  of  our  Western  arid  plains 
in  the  dry  season  the  air  is  almost  free  of  moisture. 

Character  and  Composition  of  Air  which  has  been  Breathed.— 
The  changes  effected  by  respiration  in  the  atmospheric  air  are:  i,  an  increase 
of  temperature;  2,  a  diminution  in  the  quantity  of  oxygen;  3,  an  increase  in 
the  quantity  of  carbon  dioxide;  4,  a  diminution  of  volume;  5,  an  increase  in 
the  amount  of  watery  vapor;  6,  the  addition  of  a  minute  amount  of  organic 
matter  and  of  free  ammonia. 

Temperature  of  the  Expired  Air. — Expired  air,  after  its  contact  with  the 
Interior  of  the  lungs,  is  hotter  (at  least  in  most  climates)  than  the  inspired  air. 
its  temperature  varies  between  36°  and  37.5°  C.  (97°  and  99.5°  F.),  the 
lower  temperature  being  observed  when  the  air  has  remained  but  a  short 
time  in  the  lungs.  Whatever  may  be  the  temperature  of  the  air  when  in- 
haled, it  acquires  nearly  that  of  the  blood  before  it  is  expelled  from  the  chest. 

The  Oxygen  of  Expired  Air. — Pettenkofer  and  Voit  have  found  that  the 
mean  consumption  of  oxygen  during  24  hours  by  a  man  weighing  70  kilos 
is  about  700  grams  or  490  liters.  The  quantity  of  oxygen  absorbed  increases 
with  muscular  exercise,  and  falls  during  rest.  In  general  terms  the  quantity 
absorbed  varies  with  the  activity  of  the  metabolic  processes,  following  very 
closely  the  variation  of  carbon  dioxide  under  the  conditions  outlined  below. 

The  Carbon  Dioxide  of  Expired  Air. — The  percentage  of  carbon  dioxide 
is  increased  in  expired  air,  but  the  total  quantity  of  carbon  dioxide  exhaled 
in  a  given  time  is  subject  to  change  from  various  circumstances.  From 
every  volume  of  air  inspired  4  to  5  per  cent,  of  oxygen  is  abstracted;  while 
a  rather  smaller  quantity,  4.38  per  cent.,  of  carbon  dioxide  is  added  in  its 
place.  The  expired  air  will  contain,  therefore,  438  volumes  of  carbon  di- 
oxide in  10,000.  The  total  quantity  of  carbon  dioxide  exhaled  into  the  air 
breathed  by  a  healthy  adult,  calculating  that  15.4  cc.  of  the  350  cc.  of  the 
average  air  exhaled  at  each  expiration  consists  of  carbon  dioxide,  and  that 
the  rate  of  respiration  per  minute  is  on  an  average  16,  would  be  about  400 
liters  in  twenty-four  hours.  From  actual  experiment  this  amount  seems 
to  be  a  trifle  too  great,  since  from  the  average  of  many  investigations  the 
total  amount  of  carbon  dioxide  excreted  per  day  by  the  entire  body  has  been 
found  to  be  about  400  liters,  weighing  800  grams,  and  consisting  of  218 
grams  of  carbon,  and  582  grams  of  oxygen.  From  the  218  grams  of  carbon 
must  be  deducted  about  10  grams  excreted  in  other  ways  than  by  the  lungs, 
which  leaves  about  215  grams  as  the  amount  of  carbon  excreted  by  the  aver- 
age healthy  man  by  respiration  each  day  and  night.  These  quantities 


CHARACTER    AND    COMPOSITION    OF    AIR 


299 


must  be  considered  approximate  only,  inasmuch  as  various  circumstances, 
even  in  health,  influence  the  amount  of  carbon  dioxide  excreted,  and,  cor- 
relatively,  the  amount  of  oxygen  absorbed. 

The  total  amount  of  carbon  dioxide  excreted  is  influenced  sharply  by  a 
number  of  factors:  First,  the  depth  and  volume  of  respiratory  movements.  The 
greater  the  volume  of  air  breathed,  the  greater  the  total  output  of  carbon 
dioxide,  though  the  percentage  per  unit  of 
expired  air  is  decreased.  This  influence  de- 
pends upon  the  more  efficient  oxidative 
processes  in  the  presence  of  more  thorough 
ventilation  of  the  lungs  and  blood.  Second, 
the  carbon  dioxide  output  varies  with  age. 
It  is  greater  with  children  and  youth  than 
with  the  old.  In  extreme  old  age  the  total 
output  may  not  exceed  that  of  the  ten-year- 
old  child.  Third,  there  is  a  diurnal  variation 
in  carbon  dioxide  output.  The  respiratory 
quotient,  i.e.,  the  ratio  between  carbon 
dioxide  eliminated  and  oxygen  absorbed,  is 
greater  during  the  day  than  during  the 
night.  In  the  day,  therefore,  the  carbon 
dioxide  exhaled  in  relation  to  the  oxygen 
absorbed  is  increased,  and  it  is  diminished 
during  the  night.  This  is  probably  due  to 
the  increased  production  of  carbon  dioxide 
as  a  result  of  increased  tissue  activity  during 
the  day,  and,  consequently,  the  breaking 
down  or  katabolism  of  more  substances. 
Fourth,  the  character  and  quantity  of  the 
food  greatly  influence  the  proportion  of 
carbon  dioxide  as  indicated  by  the  respira- 
tory quotient.  It  is  greater  with  carbohy- 
drate foods.  During  fasting  there  is  for  the 
first  two  or  three  days  an  increased  carbon 
dioxide  output,  but  later  this  is  decreased. 
Fifth,  the  bodily  exercise,  in  moderation, 
increases  the  quantity  of  carbon  dioxide  ex- 
pired by  at  least  one-third  more  than  it  is 

during  rest.  For  about  an  hour  after  exercise  the  volume  of  the  air  expired 
in  the  minute  is  increased  nearly  2,000  cc.,  or  118  cubic  inches;  and  the 
quantity  of  carbon  dioxide  about  125  cc.,  or  7.8  cubic  inches  per  minute. 
Violent  exercise,  such  as  full  labor  or  athletic  competition,  still  further  in- 
creases the  amount  of  the  carbon  dioxide  exhaled.  Sixth,  the  observations 
made  by  Vierordt  at  various  temperatures  between  3.4°-23.8°  C.  (38°  F. 
and  75°  F.)  show,  for  warm-blooded  animals,  that  within  this  range  every 
rise  equal  to  5.5°  C.  (10°  F.)  causes  a  diminution  of  about  33  cc.  (2  cubic 
inches)  in  the  quantity  of  carbon  dioxide  exhaled  per  minute. 

The  Volume  of  the  Respired  Air  is  Diminished. — When  allowance  has 
been  made  for  the  expansion  in  heating,  the  volume  of  expired  air  is  decreased, 


FIG.  236. — Apparatus   for  Esti- 
:>2  and  CO2  in  Expired 


300  RESPIRATION 

the  loss  being  due  to  the  fact  that  a  portion  of  the  oxygen  absorbed  is  not 
returned  in  the  form  of  carbon  dioxide.  Since  the  oxygen  of  a  given  volume 
of  carbon  dioxide  would  have  the  same  volume  as  the  carbon  dioxide  itself 
at  a  given  temperature  and  pressure,  a  portion  of  the  oxygen  absorbed 
must  be  used  for  other  purposes  than  the  formation  of  carbon  dioxide. 
In  fact,  some  of  it  is  used  in  the  formation  of  urea,  some  in  the  formation 
of  water,  etc.  The  volume  of  the  carbon  dioxide  exhaled,  divided  by  the 
volume  of  the  oxygen  absorbed,  gives  what  is  known  as  the  respiratory  quo- 
tient; thus 

CO2  exhaled 

O2  absorbed 

Normally  in  man  on  a  mixed  diet  the  respiratory  quotient  averages  0.82 

4.0  to  4.5 

=  o. 8  to  0.9. 

But  it  is  subject  to  variation  through  several  causes;  for  example,  through 
variation  in  the  composition  of  the  diet.  On  a  pure  carbohydrate  diet  the 
respiratory  quotient  will  rise  above  0.9,  i.e.,  to  i.o,  since  carbohydrates 
contain  enough  oxygen  to  oxidize  the  hydrogen  in  the  molecule.  On  a  diet 
containing  much  fat  the  quotient  is  lowest,  since  relatively  more  oxygen  is 
needed  completely  to  oxidize  fat.  The  theoretical  respiratory  quotient  for 
fats  is  0.7.  The  same  is  true,  but  to  a  less  degree,  in  the  case  of  proteins 
which  also  require  much  oxygen  for  their  complete  oxidation.  Muscular 
exertion  raises  the  respiratory  quotient,  because  in  its  performance  carbo- 
hydrates are  used  up  in  relatively  greater  quantity. 

The  Watery  Vapor  in  Respired  Air. — The  quantity  of  water  vapor  emitted 
is,  as  a  general  rule,  sufficient  to  saturate  the  expired  air,  or  very  nearly  so. 
Its  absolute  amount  is,  therefore,  influenced  by  the  following  circumstances: 
i.  By  the  quantity  of  air  respired;  for  the  greater  the  volume  of  air,  the 
greater  also  will  be  the  quantity  of  moisture  exhaled;  2.  by  the  quantity  of 
water  vapor  contained  in  the  air  previous  to  its  being  inspired;  because  the 
greater  the  moisture  inhaled,  the  less  will  be  the  amount  to  complete  the 
saturation  of  the  air;  3.  by  the  temperature  of  the  expired  air;  for  the  higher 
the  temperature  the  greater  will  be  the  quantity  of  water  vapor  required  to 
saturate  the  air;  4.  by  the  length  of  time  which  each  volume  of  inspired  air 
is  allowed  to  remain  in  the  lungs;  for  although,  during  ordinary  respiration, 
the  expired  air  is  always  saturated  with  water  vapor,  yet,  when  respiration  is 
performed  very  rapidly,  the  air  has  scarcely  time  to  be  raised  to  the  highest 
temperature  or  be  fully  charged  with  moisture  ere  it  is  expelled. 

The  quantity  of  water  exhaled  from  the  lungs  in  24  hours  ranges  (accord- 
ing to  the  various  modifying  circumstances  already  mentioned)  from  about 
200  to  800  cc.,  the  ordinary  quantity  being  about  400  to  500  cc.  Some  of 
this  is  probably  formed  by  the  chemical  combination  of  oxygen  with  hydro- 


RESPIRATORY    CHANGES    IN    THE    BLOOD  3<DI 

gen  in  the  system;  but  the  far  larger  proportion  of  it  is  water  which  has  been 
absorbed,  as  such,  into  the  blood  from  the  alimentary  canal,  and  which  is 
exhaled  from  the  surface  of  the  air-passages  and  cells,  as  it  is  from  the  free 
surfaces  of  all  moist  animal  membranes,  particularly  at  the  high  tempera- 
ture of  warm-blooded  animals. 

A  small  quantity  of  ammonia  is  added  to  the  ordinary  constituents  of 
expired  air.  It  seems  probable,  however,  both  from  the  fact  that  this  sub- 
stance cannot  be  always  detected  and  from  its  minute  amount  when  present, 
that  the  whole  of  it  may  be  derived  from  decomposing  particles  of  food  left 
in  the  mouth  or  the  teeth,  and  that  it  is,  therefore,  only  an  accidental  con- 
stituent of  expired  air. 

The  Organic  Matter  in  Expired  Air. — It  was  formerly  supposed  that  this 
organic  matter  was  injurious  and  gave  rise  to  the  unpleasant  symptoms 
which  are  experienced  in  badly  ventilated  rooms.  But  this  has  been  strongly 
questioned  so  that  the  matter  cannot  be  considered  settled  at  the  present 
time. 

THE  RESPIRATORY  CHANGES  IN  THE  BLOOD. 

Pressure  and  Diffusion  of  the  Air. — It  must  be  remembered  that 
the  tidal  air  in  the  lungs  amounts  only  to  from  300  to  500  cc.  at  each  in- 
spiration. This  amount  at  once  mixes  with  the  reserve  and  the  residual 
air  already  in  the  lungs.  The  mixture  is  facilitated  by  the  air  currents  set 
up  in  the  deeper  parts  of  the  lungs  by  the  sudden  entrance  of  the  tidal  air; 
but,  after  all  is  considered,  it  will  be  found  that  diffusion  is  the  greatest  factor 
in  producing  a  uniform  mixture  of  the  gases  in  the  alveoli  and  in  the  air-cells 
of  the  lungs.  Just  as  a  fresh  supply  of  oxygen  introduced  within  the  door 
of  a  closed  room  will  quickly  diffuse  throughout  the  space  of  the  entire  room 
so  will  the  fresh  tidal  air  diffuse  into  the  space  of  the  lungs.  When  the 
tidal  air  is  expired  its  average  composition  has  been  changed  so  it  has  only 
about  16  per  cent,  of  oxygen  instead  of  the  usual  20. 96  per  cent,  of  oxygen  in 
air.  The  oxygen  content  of  the  air  still  left  in  the  lungs  is  probably  some- 
what less  than  the  percentage  in  this  expired  air  for  the  reason  that,  the  air 
of  the  respiratory  tree,  the  trachea,  bronchi,  and  bronchioles,  is  never  fully 
mixed  with  the  alveolar  air. 

The  partial  pressure  of  the  oxygen  of  the  air  measured  under  standard 
conditions  is  159  mm.  of  mercury;  that  is,  20. 96  per  cent,  of  760  mm.  of  mer- 
cury, the  standard  pressure  of  one  atmosphere.  The  partial  oxygen  pressure 
in  expired  air  with  16  per  cent,  of  oxygen  is  only  122  mm.  of  mercury.  These 
figures  show  a  diffusion  pressure  of  at  least  37  mm.  of  mercury  to  carry 
oxygen  into  the  deeper  recesses  of  the  lungs.  The  constant  loss  of  oxygen 
to  the  blood  probably  keeps  the  mean  difference  greater. 

The  Gases  of  the  Blood. — Turning  now  to  the  consideration  of  the 
gases  of  the  blood  in  the  lungs,  a  somewhat  different  picture  presents  itself. 
19 


302 


RESPIRATION 


The  blood  consists  of  a  fluid  plasma  with  a  mass  of  corpuscles  floating  in 
it.  The  gas  analysis  of  the  blood  shows  that  it  contains  oxygen,  carbon 
dioxide,  nitrogen,  and  traces  of  other  inert  gases.  The  blood  gases  are 
measured  by  the  method  of  extracting  them,  measuring  the  volume  and 
computing  the  volume  to  standard  temperature  and  pressure. 

Numerous  analyses  of  the  blood  from  the  arteries  and  veins  of  normal 
men  have  recently  been  obtained  by  Stadie,  Harrop,  and  others,  made 
possible  by  the  development  of  the  micro-analytical  methods  and  appara- 
tus introduced  by  Van  Slyke,  Fig.  237.  Arterial  blood  obtained  by 
puncture  from  the  radial  artery  with  a  slender  hypodermic  needle  and 
syringe  have  yielded  on  analysis  the  following  average  volumes  per  cent, 
of  oxygen. 

Arterial  and  Venous  Oxygen,  Total  Oxygen  Capacity,  and  Arterial  and  Venous  Oxygen  Unsaturation 
in  Five  normal  Individuals  (Stadie) 


Oxygen  content 

Unsaturation 

Individual  number 

Oxygen 
capacity 

Arterial 

Venous 

Arterial, 

Venous, 

per  100  cc. 

per  100  cc. 

per  100  cc. 

of  blood 

of  blood 

of  blood 

Per  100  cc. 

Per 

Per  100  cc. 

Per 

of  blood 

cent. 

of  blood 

cent. 

I 

17-9 

12.8 

19.1 

I  .2 

6.3 

6.3 

33-0 

2 

21  .0 

16.7 

21.6 

0.6 

2.8 

4-9 

22.7 

3 

22.1 

17-2 

23   3 

I  .  2 

5-2 

6.1 

26.2 

4 

20.2 

15-6 

21.6 

1-4 

6.5 

6.0 

27.8 

5 

19-5 

IS-4 

20.3 

0.8 

39 

4-9 

24.1 

2O.  2 

15.6 

21  .  2 

I  .0 

S.o 

5.6 

26.8 

The  amount  of  oxygen  per  unit  quantity  of  blood  varies  with  the  con- 
centration of  hemoglobin.  In  blood  from  the  radial  artery  the  sample  is 
under  normal  respiratory  conditions  about  93  to  97  per  cent  saturated, 
See  Harrop,  Table  I. 

The  amount  of  carbon  dioxide  in  the  total  blood  averages  about  40 
volumes  per  cent,  in  arterial  blood  and  46  to  58  volumes  per  cent,  in  venous 
blood.  Venous  blood  may  contain  as  much  as  65  volumes  per  cent,  of  car- 
bon dioxide.  The  carbon  dioxide-carrying  bases  are  largely  in  the  plasma 
and  increase  or  decrease  with  variations  of  acid  or  alkali  production, 
thereby  maintaining  equilibrium.  The  amount  of  nitrogen  in  solution  in 
the  blood  follows  closely  its  ratio  of  physical  absorption  by  fluids.  Saturated 
arterial  blood  contains  1.52  volumes  per  cent,  of  nitrogen.  Venous  blood 
contains  somewhat  less,  about  1.36  volumes  per  cent.  (Van  Slyke  and 
Stadie). 


RESPIRATORY    CHANGES    IN   THE    BLOOD 


303 


Petition 


For  extracting  the  gases  from  the  blood  the  older  methods  using  the  mercurial  air 
pumps  of  Ludwig,  Geissler,  or  Sprengel  have  given  place  to  much  simpler  and  more 
convenient  microapparatus  of  Van  Slyke,  Fig.  237.  The  Van  Slyke  apparatus  can  be 
used  for  the  analysis  of  oxygen,  or  of  carbon  dioxide,  and  of  the  inert  residue  of  nitrogen 
by  difference. 

The  Van  Slyke  Blood  Gas  Apparatus.— -The  Van  Slyke  apparatus  consists  of  a  50  cc. 
pipette  with  three-way  stop  cocks,  e  and  /,  at  the  top  and  bottom.  The  top  of  the  pip- 
ette is  graduated  in  i  cc.  and  .02  cc.  divisions.  A  . 
reservoir  of  80  cc.  capacity  is  connected  with  the 
bottom  of  the  apparatus  by  a  heavy  black  rubber 
tubing  of  small  bore  and  the  whole  apparatus  filled 
with  mercury.  The  sample  of  blood  to  be  analyzed 
is  introduced  through  the  cup  b  into  the  pipette  and 
the  gases  evacuated  and  measured  according  to  the 
following  technique. 

The  solutions  required  are  ammonia  solution  to 
which  is  added  the  soluble  saponine  from  5  grams  of 
commercial  soap  bark  and  4  cc.  concentrated  am- 
monia per  liter;  redistilled  caprillic  alcohol  to  prevent  &   \mabort 
foaming;    and    10  per  cent,   potassium  ferricyanide 


Volume  of  Gas  Measured. 


Column  of  Water  Solution 

Level  of  Mercury  Surface 
in  Levelling  Bulb. 
vLevel  of  Mercury 
Menijcuj  m  Pipette. 


FIG.  2370. 


FIG.  237. 


in  normal  potassium  hydrate.  A  determination  is  made  by  the  following  steps.  Intro- 
duce 3  drops  of  caprillic  alcohol  and  6  cc.  ammonia  into  the  pipette,  evacuate  and  wash 
out  the  dissolved  air,  run  2  cc.  of  air-free  ammonia  back  up  in  the  cup  as  a  seal.  A  2  cc. 
sample  of  fresh  arterial  blood  drawn  under  oil  is  run  from  a  2  cc.  graduated  pipette  into 
the  cup  under  the  ammonia  solution  and  drawn  down  into  the  pipette.  Mix  the  solu- 
tions until  the  blood  is  completely  laked,  which  occurs  in  a  few  seconds.  Next  intro- 
duce 0.4  cc.  of  10  per  cent,  ferricyanide  to  set  the  oxygen  free  from  the  hemoglobin. 
This  dissociation  is  facilitated  by  lowering  the  mercury  to  about  the  50  cc.  mark  on  the 
pipette  thus  producing  a  Toricellian  vacuum  in  which  the  blood  and  reagent  mixture  is 


304  RESPIRATION 

shaken  vigorously  for  about  one  minute.  Make  a  preliminary  reading  of  the  liberated 
gas  and  repeat  the  evacuation  until  the  readings  check.  For  final  reading  draw  the 
fluids  into  the  chamberd  below  the  lower  stopcock,  using  care  not  to  trap  any  gas,  and 
run  the  mercury  around  the  side  tube  c,  level  the  mercury  bulb  against  the  mercury 
meniscus  in  the  graduated  limb  as  in  Fig.  2376  and  read.  The  gas  of  the  2  c.c.  sample 
of  blood  consists  of  the  oxygen  bound  by  the  hemoglobin  and  of  the  air  in  solution  at  the 
temperature  and  barometer  of  the  analysis.  The  corrections  for  dissolved  air  including 
nitrogen  are  readily  made  from  tables  of  solubility.  (See  Van  Slyke  in  Journal  of  Bio- 
logical Chemistry,  Vol.  33,  p.  126;  also  Vol.  49,  p.  i.) 

The  large  quantity  of  oxygen  found  in  arterial  and  in  venous  blood  is 
the  more  striking  when  the  facts  of  absorption  of  gases  by  liquids  are 
reviewed.  A  liquid  such  as  water  will,  when  exposed  to  a  gas,  take  up  the 
gas  by  absorption  according  to  definite  physical  laws.  Under  constant 
temperature  the  amount  of  gas  absorbed,  oxygen  for  example,  varies 
directly  as  the  pressure  of  the  gas,  or  partial  pressure  if  the  gas  is  in  a  mix- 
ture. The  oxygen  absorbed  by  water  from  pure  air  is  in  direct  pro- 
portion to  the  partial  pressure  of  oxygen  in  the  air,  which  is  159  mm. 
mercury. 

The  amount  of  gas  absorbed  by  i  c.c.  of  water  under  standard  pressure 
(one  atmosphere  at  oc  C.)  is  termed  the  absorption  coefficient.  The 
absorption  of  oxygen  by  water  for  one  atmosphere  of  oxygen  is  .048  c.c. 
For  blood  plasma  the  coefficient  is  a  little  less  than  for  water.  The 
amount  of  oxygen  in  simple  solution  in  100  c.c.  of  blood  at  the  partial 
pressure  of  oxygen  in  alveolar  air  is  therefore  only  about  0.32  c.c.  The 
actual  amount  of  oxygen  in  solution  in  any  particular  specimen  of  plasma 
is  rather  less  and  is  determined  by  the  oxygen  tension. 

The  saturation  of  oxygen  in  arterial  whole  blood  is  measured  by  the 
method  of  subjecting  the  blood  to  an  atmosphere  in  which  the  oxygen 
tension  is  accurately  known.  The  instrument  is  called  a  tonometer. 
The  procedure  depends  upon  the  fact  that  a  thin  film  of  blood  exposed  to 
mixtures  of  gases  in  air  gives  up  gases  to  or  absorbs  them  from  the  air 
until  an  equilibrium  is  established.  When  a  sample  of  whole  blood  is 
exposed  to  atmospheric  air  in  a  tonometer  the  blood  becomes  fully  satur- 
ated with  oxygen  and  the  volume  it  contains  is  spoken  of  as  the  capacity. 
When  such  blood  has  its  gases  extracted  by  the  Van  Slyke  apparatus  and 
the  results  computed  to  standard,  the  volumes  per  cent,  contained  are 
such  as  indicated  in  the  table,  page  302.  When  alveolar  airs  are  used 
the  degree  of  saturation  is  of  course  proportionately  less  than  the  satu- 
ration against  pure  air  because  of  the  diminished  per  cent,  of  alveolar 
oxygen.  The  volumes  per  cent,  of  oxygen  absorbed  is  found  to  vary  also 
according  to  the  per  cent,  of  oxygen  in  the  sample  of  air  and  the  content 
of  hemoglobin  in  the  blood. 

By  means  of  the  tonometer  observers  have  measured  the  tension  of 


RESPIRATORY    CHANGES    IN    THE   BLOOD 


305 


blood  gases.  The  oxygen  tension  has  been  found  to  be  from  4  (S trass- 
burg)  to  10  (Herter)  per  cent,  of  an  atmosphere.  Many  determinations 
have  been  given  of  both  lower  and  higher  percentages,  but,  accepting  the 
above  limits  for  a  working  average,  the  oxygen  tension  in  arterial  blood 
would  be  from  30.4  to  76  mm.  of  mercury  or  more. 

Blood  plasma  exposed  to  an  air  with  a  partial  pressure  of  30  to  76  mm. 
of  mercury  would  absorb  only  from  o.io  to  0.32  (0.26  c.c.  Pfluger)  c.c. 
of  oxygen  for  100  c.c.  of  blood.  As  a  matter  of  fact  100  c.c.  of  whole 


100 
90 

80 
70 
60 
50 
40 
30 
20 
10 


0  10         20         30         40         50         60         70         80         90      100 

FIG.  238. — Dissociation  curves  of  oxyhemoglobin.  The  figures  along  the  ordinates 
represent  percentages  of  saturation  of  hemoglobin  by  oxygen.  The  figures  along  the 
abscissae  represent  mm.  of  oxygen  pressure  in  mercury. 

I.  Bohr's  dissociation  curve  of  oxyhemoglobin  dissolved  in  water. 

II.  Dissociation  curve  of  oxyhemoglobin  dissolved  in   Ringer's  Solution.     (After 
Barcroft  and  Camis.) 

blood  has  a  capacity  of  an  average  of  from  18.5  c.c.  (the  Haldane  stand- 
ard) to  22.6  c.c.  or  more  of  oxygen.  It  is  evident  that  blood  carries  far 
more  oxygen  than  can  be  held  in  simple  solution.  The  red  blood  corpus- 
cles carry  their  enormous  excess  of  oxygen  by  virtue  of  the  special  respira- 
tory pigment,  hemoglobin. 


3o6 


RESPIRATION 


Combining  Power  of  Hemoglobin  with  Oxygen. — One  hundred 
cubic  centimeters  of  blood  contain  about  14  grams  of  hemoglobin,  page 
137.  Each  gram  of  hemoglobin,  when  fully  saturated  with  oxygen,  accord- 
ing to  Hiifner's  earlier  determination,  combines  with  1.56  cc.  of  oxygen. 
By  later  work  he  gets  the  determination  of  1.34  cc.  for  hemoglobin  of  ox 
blood.  This  last  figure  indicates  that  the  combining  power  of  the  hemo- 
globin is  dependent  upon  the  iron  in  the  molecule,  in  which  one  atom  of  iron 
combines  with  one  atom  of  oxygen.  A  number  of  investigators  have 


too 


80 


70 


50 


40 


30 


20 


10 


10 


20 


30 


40 


5<T 


60 


70 


80 


90 


100 


FIG.  239. — Dissociation  curves  of  oxy hemoglobin  in:  I,  0.7  per  cent,  sodium  chlor- 
ide; II,  in  sodium  bicarbonate,  and  III,  in  disodium  phosphate.  The  figures  along  the 
ordinates  represent  percentages  of  saturation  of  hemoglobin  by  oxygen.  The  figures 
along  the  abscissae  represent  mm.  of  oxygen  pressure  in  mercury.  (Barcroft  and  Camis.) 

examined  the  conditions  under  which  hemoglobin  combines  with  oxygen — 
Hiifner,  Bohr,  Lowy,  and  Barcroft  and  Camis.  Hiifner,  working  with 
purified  hemoglobin  in  watery  solution,  found  that  when  the  oxygen  ten- 
sion in  the  air  in  contact  with  the  hemoglobin  was  increased  above  zero  by 
graded  stages,  the  amount  of  oxygen  that  was  combined  was  very  great  per 
unit  of  increased  pressure  at  the  low  pressures,  but  relatively  less  at  the 


POWER    OF   HEMOGLOBIN   WITH    OXYGEN 


307 


higher  pressures.  Or,  which  amounts  to  the  same,  if  hemoglobin  saturated 
with  oxygen  be  subjected  to  decreasing  oxygen  pressure,  it  sets  free  the 
combined  oxygen,  at  first  slowly,  then  more  rapidly.  By  consulting  the 
typical  curve  showing  this  relation,  it  will  be  evident  that  the  critical 
partial  oxygen  pressures  influencing  this  combination  fall  at  about  30  to 
35  mm.  mercury  of  oxygen  tension  and  below.  See  figure  238. 

A  number  of  factors  influence  the  dissociation  of  oxygen  from  hemo- 
globin at  a  given  oxygen  tension.  Of  prime  importance  is  the  influence  of 
the  presence  of  carbon  dioxide  gas  as  shown  by  Bohr  and  confirmed  by 
Barcroft  and  Camis.  With  an  increase  in  the  tension  of  carbon  dioxide 
there  is  a  decrease  in  the  fixation  of  oxygen. 

TABLE  SHOWING  THE  DISSOCIATION  OF  OXHEMOGLOBIN  IN  THE  PRESENCE  OF  VARY- 
ING TENSIONS  OF  CARBON  DIOXIDE.     (BARCROFT  AND  CAMIS.) 


Tensions  of  oxygen  in  mm.  mercury  and  the  per  cent,   of    saturation 

Tension  of  car- 

of the  hemaglobin  at  each  pressure 

bon  dioxide 

10 

IS 

20 

25 

3° 

35 

40 

45 

50 

60 

70 

80 

100 

5  mm.  COz 

28 

35 

47 

58.5 

70 

81 

89 

93 

95 

96 

98 

99 

99-5 

10  mm.  COz 

ii 

26 

38.5 

Si 

63 

74-5 

83 

88 

9I.S 

94-5 

96.5 

97.5 

98.5 

20  mm;  CO2 

0 

10 

25 

49 

S3-  8 

65.5 

74-5 

80.5 

84.5 

90 

93 

95 

97-5 

40  mm.  COz 

0 

o 

1  1 

26 

42,5 

56 

65 

72 

77 

83.5 

88.  5 

93 

95-5 

80  mm.  COz 

0 

o 

I 

12-5 

3i 

45-5 

56.5 

64 

69.5 

77 

83 

87.5 

92.5 

The  salts  of  the  blood  also  influence  the  oxygen  fixation  by  hemo- 
globin under  a  given  tension  as  indicated  in  the  following  table: 

TABLE  SHOWING  THE  INFLUENCE  OF  THE  PRESENCE  OF  DIFFERENT  SALTS  ON 

THE  PERCENTAGE  OF  SATURATION  OF  HEMOGLOBIN  UNDER  A 

CONSTANT  OXYGEN  TENSION  OF  30  MM.  MERCURY. 

(BARCROFT  AND  CAMIS.) 

1.  Hemoglobin  in  water  dissociation . : 62  per  cent. 

2.  Hemoglobin  in  o .  7  per  cent.  NaCl  dissociation ....    75  per  cent. 

3.  Hemoglobin  in  Ringer's  solution  dissociation 85  per  cent. 

4.  Hemoglobin  in  NaHCO3  solution  dissociation 89  per  cent. 

5.  Hemoglobin  in  0.9  per  cent.  KC1  dissociation 91  per  cent. 

6.  Hemoglobin  in  Na2HPO4  solution  dissociation.  ...    93  per  cent. 

Barcroft  and  Camis  find  that  the  dissociation  curve  also  varies  in  the 
blood  of  different  animals.  Strassburg  gives  the  oxygen  tension  of  arterial 
blood  as  29.64  mm.  of  mercury,  and  for  venous  blood  22.04  mm.  of  mer- 
cury. That  is  to  say,  during  the  brief  interval  in  which  the  blood  is  in 
the  pulmonary  capillaries  the  oxygen  tension  has  increased  by  7.6  mm. 
of  mercury,  an  increase  of  tension  which  would  produce  very  little  increase 
in  simple  absorption  of  oxygen.  Yet  it  is  sufficient  to  cause  fixation  of 
from  four  to  five  volumes  per  cent,  of  oxygen  by  the  hemoglobin. 

It  is  evident  that  there  will  be  diffusion  of  oxygen  from  the  high  tension 


308  RESPIRATION 

toward  the  lower  and  in  the  direction  indicated  by  the  arrows  in  the  table 
below.  As  fast  as  the  oxygen  diffuses  into  the  venous  blood,  thus  tending  to 
raise  the  pressure  of  the  gas  in  solution,  it  is  taken  up  and  fixed  by  the  hemo- 
globin. This  process  proceeds  far  enough  during  the  interval  the  blood  is 
in  the  pulmonary  capillaries  to  raise  the  oxygen  tension  from  22.04  mm. 
of  mercury  to  29 . 64  mm.  of  mercury,  and  also  far  enough  to  permit  of  the 
fixation  of  from  four  to  five  volumes  per  cent,  of  oxygen.  The  oxygen 
diffusion  pressures  are  indicated  as  follows: 

Oxygen  pressure  in  the  atmosphere  2 1  per  cent,  or  1 59       mm.  of  mercury 

I 

Oxygen  pressure  in  the  alveolar  air  16  per  cent,  or  122       mm.  of  mercury 

1 

Oxygen  pressure  in  the  venous  blood  3  per  cent,  or  22  .04  mm.  of  mercury 

Liberation  of  Oxygen  in  the  Tissue  Capillaries. — When  the  arterial 
blood  reaches  the  capillaries  of  the  tissues,  then  the  situation  which  we  have 
just  found  holding  good  in  the  lungs  is  reversed.  As  rapidly  as  the  oxygen 
reaches  the  living  protoplasm  of  the  tissues  it  enters  into  fixed  combination, 
thus  rendering  it  inert.  The  oxygen  tension  in  the  tissue  cells  will,  there- 
fore, be  zero.  Under  these  conditions  the  difference  in  pressure  level  be- 
tween the  oxygen  tension  in  the  blood  and  that  in  the  tissues  is  sufficient  to 
cause  a  rapid  diffusion  of  oxygen  through  the  capillary  walls  with  correspond- 
ing liberation  of  the  oxygen  from  the  hemoglobin  according  to  the  laws  of 
combination  given  in  the  curves  above.  The  total  effect  of  this  process  is  to 
maintain  a  relatively  high  and  constant  diffusion  pressure  of  the  oxygen  in 
the  blood.  During  the  time  the  blood  remains  in  the  capillaries  the  total 
oxygen  tension  will  have  been  lowered  from  29 .64  to  22  .04  mm.  of  mercury, 
yet  this  slight  lowering  of  tension  is  sufficient  to  liberate  from  four  to  five 
volumes  per  cent  of  oxygen.  This  figure,  of  course,  is  comparative.  In 
many  of  the  very  active  tissues,  such  as  in  muscle,  a  much  larger  per  cent, 
of  oxygen  will  have  been  dissociated  and  the  oxygen  tension  correspondingly 
lowered  so  that  the  venous  blood  returning  through  such  an  active  organ 
may  not  have  more  than  half  the  average  amount  of  oxygen  found  in  venous 
blood. 

Considering  the  pressure  relations  of  oxygen  from  the  time  of  its  intro- 
duction into  the  body  with  the  fresh  air  to  its  fixation  in  the  tissues  we  have 
the  following  schema: 

Oxygen  pressure  in  the  atmosphere 1 59      mm. 

I 

Oxygen  pressure  in  the  alveolar  air 122      mm. 

I 

Oxygen  pressure  in  the  venous  blood 22  .04   mm. 

Tension  of  oxygen  in  the  arterial  blood 29 . 64   mm. 

! 

Tension  of  oxygen  in  the  tissues o .  oo  mm. 


GASES    IN   THE    LUNGS    AND    TISSUES  309 

Elimination  of  Carbon  Dioxide  by  the  Blood  and  the  Respiratory 
Apparatus. — The  principles  of  absorption  of  gas  by  liquids  discussed 
in  the  preceding  pages  apply  equally  well  for  carbon  dioxide  with  the  excep- 
tion that  carbon  dioxide  is  about  three  times  as  soluble  in  blood  as  is  oxygen. 
The  carbon  dioxide  results  from  the  oxidative  processes  going  on  in  the  tis- 
sues, and  this  gas  is  present  in  large  quantities  in  the  tissues  and  their  im- 
mediately surrounding  lymph.  An  analysis  of  the  carbon- dioxide  content 
of  venous  blood  reveals  the  presence  of  about  45  cc.  of  the  gas  in  100  cc.  of 
blood.  This  gas,  like  oxygen,  is  held  in  such  large  quantity  by  virtue  of  the 
fact  that  it  forms  loose  chemical  combinations  in  the  blood.  Of  the  total 
quantity  not  more  than  5  per  cent,  is  held  in  simple  solution.  From  10  to 
15  per  cent,  of  the  total  volume  is  found  in  firm  combination  in  such  forms 
as  carbonates,  bicarbonates,  etc.  The  remaining  85  and  more  volumes 
per  cent,  is  held  in  loose  chemical  combination,  a  combination  which  is  broken 
up  under  the  same  conditions  of  variation  in  carbon-dioxide  tension  as 
were  found  to  exist  for  oxygen  in  combination  with  hemoglobin.  In  the  case 
of  carbon  dioxide  an  analysis  of  plasma  reveals  the  fact  that  the  gas  is  in  com- 
bination with  some  compound  of  the  plasma,  probably  a  protein.  In  fact, 
there  is  some  evidence  to  show  that  carbon  dioxide  combines  with  the  globu- 
lin group.  Carbon  dioxide  also  forms  loose  chemical  compounds  with  the 
constituents  of  the  red  corpuscles,  probably  with  the  protein  portion  of  the 
hemoglobin  molecule.  The  pressure  relations  of  this  gas  as  regards  its 
diffusion  in  the  process  of  elimination  are  shown  in  the  following  table: 

Carbon-dioxide  tension  in  the  tissues 58  mm.  of  mercury 

! 

Carbon-dioxide  tension  in  the  venous  blood  45  mm.  of  mercury 

1 

Carbon-dioxide  tension  in  the  alveolar  air .  .    23  to  38  mm.  of  mercury 

I 

Carbon-dioxide  tension  in  the  expired  air ...    5.8   mm.    of  mercury 

Theories  of  Interchange  of  Gases  in  the  Lungs  and  in  the  Tissues.— 
The  above  discussion  is  on  the  basis  of  the  mechanical  interpretation  of  the 
transfer  of  gases  in  the  lungs  and  in  the  tissues.  By  this  theory  it  is  assumed 
that  the  oxygen  passes  from  the  air  in  the  lungs  through  the  moist  pulmonary 
membrane  of  the  alveoli  through  the  capillary  walls  and  into  the  blood 
plasma,  obeying  the  physical  laws  of  gas  diffusion.  Likewise  in  the  tissues 
this  theory  presupposes  that  the  difference  in  the  mechanical  tension  in  the 
capillary  blood  plasma,  the  lymph,  and  the  living  tissue  will  lead  to  diffusion 
of  the  oxygen  in  the  direction  of  lowest  pressure,  i.e.,  toward  the  tissues. 

Some  facts  have  indicated  that  we  cannot  account  for  the  transference 
of  oxygen  by  the  purely  mechanical  theory.  The  idea  has  been  advanced 
that  the  living  epithelial  wall  of  the  lung,  as  well  as  that  of  the  capillaries, 
exerts  a  distinct  influence  on  the  passage  of  oxygen  of  such  nature  that  it 


310  RESPIRATION 

might  be  regarded  as  a  secretion  of  this  gas.  This  theory  finds  some 
support  in  that  a  distinct  secretion  of  oxygen  in  the  air  bladders  of  certain 
fishes  has  been  proven  by  Bohr.  The  theory  apparently  does  not  apply 
to  mammals. 

THE  NERVOUS  REGULATION  OF  THE  RESPIRATORY 
APPARATUS. 

Respiratory  movement  is  essentially  an  involuntary  act.  Unless  this 
were  the  case,  life  would  be  in  constant  danger,  and  would  cease  on  the 
loss  of  conciousness  for  a  few  moments,  as  in  sleep.  It  is,  however,  of  ad- 
vantage to  the  body  that  co-ordination  of  respiratory  movements  should 
be  to  some  extent  under  the  control  of  the  will.  For,  were  it  not  so,  it 
would  be  impossible  to  perform  those  voluntary  respiratory  acts,  such 
as  speaking,  singing,  and  the  like. 

The  Respiratory  Nerve  Center.  —  It  has  been  known  for  centuries 
that  there  exists  a  region  of  the  central  nervous  system  on  the  destruction  of 
which  both  respiration  and  life  cease.  Flourens,  1842,  after  many  series 
of  experiments  as  to  the  exact  position  of  what  he  called  the  "knot  of  life" 
(n&ud  -vital) ,  placed  it  in  the  floor  of  the  fourth  ventricle,  at  the  point  of  the  V 
in  the  gray  matter  at  the  lower  end  of  the  calamus  scriptorius;  a  district  of 
considerable  size,  some  5  mm.  in  extent  on  each  side  of  the  middle  line. 
Observers  subsequent  to  Flourens  have  attempted  to  show  that  the  chief 
respiratory  center,  on  the  one  hand,  is  situated  higher  up  in  the  nervous 
system,  in  the  floor  of  the  third  ventricle  (Christiani),  or  in  the  corpora  quad- 
rigemina  (Martin  and  Booker,  Christiani,  and  Stanier),  or  lower  down  in 
the  spinal  cord.  The  balance  of  experimental  evidence,  however,  is  to  prove 
that  the  sole  centers  for  respiration  are  in  a  limited  district  in  the  medulla 
oblongata  in  close  connection  with  the  vagus  nucleus  on  each  side.  They 
are  approximately  identical  in  location.  The  destruction  of  this  region  stops 
respiration.  If  the  center  be  left  in  connection  with  the  muscles  of  respira- 
tion by  their  nerves,  although  the  remainder  of  the  central  nervous  system  be 
separated  from  it,  respiration  continues.  It  may  be  considered  almost  cer- 
tain that  the  medullary  center  is  the  only  true  respiratory  center.  Langen- 
dorff  states  that  in  newly  born  animals  in  which  the  medulla  has  been  im- 
mediately cut  across  at  a  level  a  few  millimeters  below  the  point  of  the 
calamus  scriptorius,  respiration  continues  for  some  time,  but  this  is  ques- 
tionable. Normal  respiration  does  not  occur  after  separation  of  the  bulb 
from  the  cord,  and  the  so-called  respiratory  movements  noticed  by  Langen- 
dorff  are  merely  tetanic  contractions  of  the  respiratory  muscles  in  which  often 
enough  other  muscles  take  part. 

The  action  of  the  medullary  center  is  to  send  out  impulses  during  in- 
spiration, which  cause  contractions  of  the  inspiratory  muscles — a,  of  the 
nostrils  and  jaws,  through  the  facial  and  inferior  division  of  the  fifth  nerves; 
b,  of  the  glottis,  chiefly  through  the  inferior  laryngeal  branches  of  the  vagi;  c 


ACTION    OF    AFFERENT    STIMULI    ON   THE    RESPIRATORY    RHYTHM      311 


of  the  intercostal  and  other  muscles  which  produce  raising  of  the  ribs,  chiefly 
through  the  intercostal  nerves,  and  d,  of  the  diaphragm,  through  the  phrenic 
nerves.  If  any  one  of  these  sets  of  nerves  be  divided,  respiratory  movements 
of  the  corresponding  muscles  cease.  Similarly  it  may  be  supposed  that 
the  center  sends  out  impulses  to  certain  other  muscles  during  expiration. 

It  has  been  suggested,  however,  that  the  center  is  double,  that  it  is  made 
up  of  inspiratory  cells  which  are  constantly  in  action,  and  of  an  expiratory 
group  of  cells  which  act  less  generally,  inasmuch  as  ordinary  tranquil  ex- 
piration is  seldom  more  than  an  elastic  recoil,  and  not  a  muscular  act  to 
any  marked  degree. 

The  respiratory  center  is  also  bilateral,  as  has  been  proven  by  the  method 
of  antero-posterior  section  of  the  medulla.  The  tracts  from  each  half  of  the 
center  are  separate  and  distinct.  If  the  cervical  cord  be  split  into  a  right 
and  left  half,  and  one  side  sectioned  at  the  level  of  the  second  cervical  ver- 
tebra, then  the  respiratory  movements  of  that  side  of  the  diaphragm  cease 
while  on  the  opposide  side  they  continue  their  rhythm. 

Assuming  this  view  of  the  quadruple  nature  of  the  respiratory  centers 
to  be  correct,  there  is  some  difference  of  opinion  as  to  the  exact  working 
of  the  mechanism  in  its  reactions.  It  is  thought  that  the  center  may  act 
automatically,  but  normally  its  automatic  discharges  of  nerve  impulses  are 


p° 

If?         R.  96  per 


BOTH  VAGI  CUT 


JYW 


•'^ 

R.  120 


FIG.  240. — The  Effect  on  the  Respiratory  Rate  of  Cutting  Both  Vagi  in  the  Dog. 
The  rate  of  60  respirations  per  minute  before  the  section  of  the  nerves  drops  to  8  per 
minute  afterward.  The  arterial  blood-pressure  is  also  shown,  the  pressure  in  mm.  mer- 
cury is  shown  in  the  scale  to  the  left.  Compare  with  figure  182. 

influenced  by  afferent  impulses  from  the  periphery,  as  well  as  by  impulses 
passing  down  from  the  cerebrum.  The  center  is,  in  other  words,  both 
automatic  and  reflex.  It  will  be  simplest  to  discuss  its  reflex  function  first. 
Action  of  Afferent  Stimuli  on  the  Respiratory  Rhythm. — Action 
of  the  vagi.  If  both  vagi  be  divided  in  the  neck,  the  respirations  become 
much  slower  and  deeper,  figure  240.  This  may  be  the  case,  but  to  a  less 
marked  degree,  if  one  of  the  nerves  is  divided  instead  of  both.  If  the  cen- 
tral, end  of  the  divided  nerve  be  stimulated  with  a  weak  but  properly  adjusted 


312  RESPIRATION 

strength  of  interrupted  current,  the  effect  is  to  quicken  the  respirations. 
And  if  the  stimuli  are  properly  regulated  the  normal  rhythm  of  respira- 
tion may  be  approached.  If  the  stimuli  be  repeated  with  stronger  currents, 
the  breathing  is  brought  to  a  standstill,  sometimes  at  the  height  of  inspira- 
tion, by  tetanus  of  the  diaphragm.  Usually,  however,  stimulation  of  the 
central  end  of  the  divided  vagus  produces  still  greater  slowing  than  that 
which  follows  the  division,  so  that  the  respirations  cease  with  the  diaphragm 
in  a  condition  of  complete  relaxation,  figures  205  and  241. 

The  sensory  action  of  the  vagus  may  therefore  be  to  call  forth  either 
inspiration  or  expiration — the  impulses  passing  up  the  vagi  being  factors 
for  the  production  and  regulation  of  the  normal  variations  in  respiratory 
rhythm.  The  fibers  of  the  vagus  are  stimulated  under  the  following  cir- 
cumstances: one  set  of  fibers,  those  which  tend  to  inhibit  expiration  and 
to  stimulate  inspiration,  are  stimulated  at  their  origin  in  the  lung  when  the 
lung  tissue  is  under  least  tension,  i.e.,  in  a  condition  of  expiration.  The 
fibers  which  tend  to  inhibit  inspiration  and  to  promote  expiration  are  stim- 
ulated when  the  lung  is  fully  expanded.  The  afferent  impulses  by  this  view 
are  the  results  of  mechanical  stimulation,  and  do  not  depend  upon  the 
chemical  nature  of  the  gases  within  the  pulmonary  alveoli. 


FIG.  241. — The  Effect  of  Stimulating  the  Vagus  on  Respiratory  Rate.  The  stimulus 
was  applied  between  the  points  "on"  and  "off."  The  inhibition  lasts  some  seconds  after 
the  stimulus  is  removed.  Time  in  seconds.  The  intra-tracheal  pressure  is  recorded 

The  Respiratory  Action  of  the  Superior  Laryngeal  Nerves. — If  the 
superior  laryngeal  branch  of  the  vagus  be  divided,  which  usually  produces 
no  apparent  effect,  and  the  central  end  be  stimulated,  the  reaction  is  very 
constant,  respirations  are  slowed,  and  there  is  a  distinct  tendency  toward 
expiration,  as  shown  by  the  contractions  of  the  abdominal  muscles.  Thus 
the  superior  laryngeal  fibers  inhibit  inspiration  and  stimulate  expiration, 
while  the  deep  branch  of  the  vagus  contains  fibers  which  stimulate  inspiration 
and  inhibit  expiration. 

The  superior  laryngeal  nerves  are  true  expiratory  nerves,  and  are  nor- 
mally set  in  action  when  the  mucous  membrane  of  the  larynx  is  irritated. 
They  are  not  in  constant  action  like  the  vagi. 

Action  of  the  Glosso-pharyngeal  Nerves. — It  has  been  ascertained, 
by  the  researches  of  Marckwald,  that  while  division  of  the  glosso-pharyngeal 
nerves  produces  no  effect  upon  respiration,  stimulation  causes  inhibition  of 
inspiration  for  a  short  period.  This  action  accounts  for  the  very  necessary 


AUTOMATIC   ACTION    OF    THE    RESPIRATORY   CENTERS  313 

cessation  of  breathing  during  swallowing.  The  effect  of  the  stimulation  is 
only  temporary,  and  is  followed  by  normal  breathing  movements. 

Action  of  Other  Sensory  Nerves. — The  respiratory  center  is  in- 
fluenced strongly  by  afferent  nerve  impulses  having  their  origin  in  general 
sensory  nerves,  particularly  the  nerves  of  the  skin.  Cold  water  suddenly 
dashed  on  the  skin  is  followed  by  a  deep  inspiration.  Stimulation  of  the 
splanchnics  or  of  the  abdominal  branches  of  the  vagi  produces  expiration. 
Stimulation  of  the  isolated  sciatic  nerve  of  the  dog  or  of  the  rabbit  causes  a 
marked  acceleration  both  of  the  rate  and  of  the  amplitude  of  the  respiratory 
movements,  see  figure  246  b.  This  acceleration  is  due  to  afferent  impulses 
which  reach  the  respiratory  center  in  the  medulla  over  sensory  paths,  paths 
which  are  not  necessarily  special  respiratory  afferent  paths,  but  rather  are 
general  afferent  paths  which  affect  the  respiratory  center  through  their 
numerous  collaterals  in  the  brain  stem. 

It  must  be  remembered  that,  although  many  sensory  nerves  may  on 
stimulation  be  made  to  produce  an  effect  upon  the  respiratory  center,  yet 
there  is  no  evidence  to  show  that  any  one  of  them,  except  the  vagus,  is  con- 
stantly in  action.  The  vagi  indeed  are,  as  far  as  we  know,  the  normal 
regulators  of  respiratory  movements,  yet  it  is  possible  reflexly  to  influence 
the  respiratory  rate  and  depth  through  impulses  that  may  have  their  origin 
in  any  sensory  part  of  the  body. 

The  respiratory  center  is  also  influenced  by  nerve  activity  of  the  cerebral 
cortex,  psychic  activity.  This  is  illustrated  by  the  limited  voluntary  control 
of  the  respiratory  movements. 

Automatic  Action  of  the  Respiratory  Centers.— Although  it  has 
been  very  definitely  proved  that  the  respiratory  centers  may  be  affected  by 
afferent  stimuli,  and  particularly  by  those  reaching  them  through  the  vagi, 
there  is  reason  for  believing  that  the  center  is  capable  of  sending  out 
efferent  nerve  impulses  to  the  respiratory  muscles  without  the  action  of  any 
afferent  stimuli.  Thus,  if  the  brain  be  removed  above  the  bulb,  respiration 
continues.  If  the  spinal  cord  be  divided  immediately  below  the  bulb,  the 
facial  and  laryngeal  respiratory  movements  continue,  although  no  afferent 
impulses  can  reach  the  center  except  through  the  cranial  sensory  nerves,  and 
these  indeed  may  be  divided  without  producing  any  effect,  when  the  bulb  and 
cord  are  intact.  As  has  been  shown,  too,  respiration  continues  when  the  vagi 
are  divided.  Isolation  of  the  respiratory  center  from  its  sensory  relations 
does  not  destroy  respiratory  movements  so  long  as  the  motor  paths  through 
the  phrenic  nerves  are  intact.  All  of  these  experiments  render  it  highly 
probable  that  afferent  impulses  are  not  required  in  order  that  the  respiratory 
centers  should  send  out  efferent  impulses  to  the  respiratory  muscles.  The 
center,  then,  is  automatic. 

Method  of  Automatic  Stimulation  of  the  Respiratory  Center. — The 
respiratory  center  is  capable  of  working  automatically  apart  from  afferent 


314  RESPIRATION 

impulses,  and  this  fact  has  been  explained  by  the  supposition  that  it  is 
.stimulated  to  action  by  the  condition  of  the  blood  circulating  through  it. 
When  the  blood  becomes  more  and  more  venous  the  action  of  the  center 
becomes  more  and  more  energetic,  and  if  the  air  is  prevented  from  entering 
the  chest,  the  respiration  in  a  short  time  becomes  very  labored.  If  the 
aeration  of  the  blood  is  much  interfered  with,  not  only  are  the  ordinary 
respiratory  muscles  employed,  but  also  those  muscles  of  extraordinary  in- 
spiration and  expiration  which  have  been  previously  enumerated.  Thus, 
as  the  blood  becomes  more  and  more  venous,  and  by  venous  we  mean  that 
the  blood  contains  a  relatively  large  amount  of  carbon  dioxide  and  a  dimin- 
ished amount  of  oxygen,  the  action  of  the  medullary  center  becomes  more 
and  more  profound.  The  question  has  been  much  debated  as  to  what 
quality  of  the  venous  blood  it  is  which  causes  this  increased  activity;  whether 
it  is  its  deficiency  of  oxygen  or  its  excess  of  carbon  dioxide.  It  has  been 
answered  to  some  extent  by  experiments  which  offer  no  obstruction  to  the  exit 
of  carbon  dioxide,  as  when  an  animal  is  placed  in  an  atmosphere  of  nitrogen. 
Under  these  conditions  dyspnea  occurs,  showing  that  blood  which  contains 
a  diminished  amount  of  oxygen  stimulates  the  cells  of  the  respiratory  center. 
On  the  other  hand,  if  the  normal  amount  of  oxygen  is  supplied  while  the 
carbon  dioxide  of  the  blood  is  prevented  from  escaping  and  thus  allowed  to 
accumulate  in  the  blood,  there  is  also  a  great  increase  in  the  respiratory 
activity  of  the  center;  an  excess  of  carbon  dioxide  in  the  blood,  flowing 
through  the  respiratory  center,  stimulates  the  cells  to  greater  activity.  It 
is  highly  probable,  therefore,  that  the  respiratory  centers  may  be  stimulated 
to  action  both  by  the  absence  of  sufficient  oxygen  in  the  blood  circulating 
in  it,  and  by  the  presence  of  an  excess  of  carbon  dioxide. 

These  facts  are  particularly  well  supported  by  the  experiments  of  Zuntz 
who  varied  the  oxygen  and  the  carbon-dioxide  content  of  the  air  breathed, 
and  measured  the  volume  breathed  per  minute.  When  the  oxygen  of  the 
air  breathed  was  reduced  by  10  to  50  per  cent.,  the  air  breathed  was  increased 
only  slightly,  5  to  10  per  cent.  When  the  oxygen  of  the  air  was  reduced 
by  60  per  cent.,  the  volume  of  air  breathed  was  increased  30  to  40  per  cent., 
and  even  more.  Other  observations  show  us  that  the  oxygen  in  the  blood 
in  these  experiments  will  fall  in  much  less  per  cent,  than  the  reduction  in 
the  oxygen  of  the  air  would  lead  us  to  suspect. 

When  Zuntz  kept  the  oxygen  content  of  the  air  about  constant,  but  in- 
creased the  carbon-dioxide  content,  then  the  amount  of  air  breathed  was 
greatly  increased.  Air  containing  18.4  per  cent,  of  oxygen  and  11.5  per  cent, 
of  carbon  dioxide  caused  an  increase  in  the  amount  breathed  per  minute 
from  7.5  liters  to  32.5  liters.  These  experiments  indicate  that  within  the 
limits  of  its  normal  variations  in  blood  the  carbon  dioxide  has  a  much  greater 
influence  than  oxygen  on  the  irritability  of  the  cells  of  the  respiratory  center. 

But  this  is  not  all,  since  it  has  been  observed  by  Marckwald  that  the 


TYPES    OF   RESPIRATION  315 

medullary  center  is  capable  of  acting  for  some  time  in  the  absence  of  any 
circulation  and  after  excessive  bleeding.  The  view  taken  by  this  author 
with  regard  to  the  action  of  the  center  is  as  follows:  The  respiratory  center 
is  set  to  act  by  the  condition  of  its  metabolism,  much  in  the  same  way  as 
the  heart  is  set  to  beat  rhythmically.  When  anabolism  is  completed,  katab- 
olism  or  discharge  occurs,  and  this  alternate  but  crude  and  spasmodic 
action  will  occur  without  a  definite  blood  supply  so  long  as  the  centers  are 
properly  nourished  and  stimulated  by  their  own  intercellular  fluid. 

It  is  also  apparent  that  the  respiratory  center  is  dependent  on  the 
character  of  the  blood  supply,  both  as  regards  quantity  and  quality  of  the 
blood.  It  has  been  shown  that  the  presence  in  the  blood  of  the  products 
of  vigorous  muscular  metabolism  will  greatly  increase  the  irritability  of  the 
respiratory  center,  even  if  the  blood  itself  be  not  particularly  venous  in 
character  as  regards  its  gaseous  content. 

The  Establishment  of  Respiratory  Movements  at  Birth. — From 
the  preceding  paragraph  it  appears  that  the  regulation  of  the  respiratory 
movements  is  normally  due  to  the  automaticity  of  the  respiratory  center  as 
influenced,  first,  by  the  blood  flowing  through  it  and,  second,  by  the  afferent 
nerve  impulses  which  reach  the  center.  The  fetus  in  the  womb  is  supplied 
by  arterial  blood  from  the  blood  vessels  of  the  mother.  The  fetus  does  not 
ordinarily  give  respiratory  movements  before  birth,  but  it  may  be  made  to 
do  so  by  experimental  procedure.  At  birth  the  placental  circulation  is  sud- 
denly interrupted,  and  the  blood  rapidly  increases  in  venosity  until  the  skin, 
lips,  and  mucous  membranes  are  very  cyanotic  in  appearance.  It  is  at  this 
time  that  the  respiratory  center  begins  its  rhythmic  discharges,  being  aroused 
by  the  direct  stimulating  effects  of  the  great  excess  of  carbon  dioxide  in  the 
strongly  venous  blood.  The  irritability  of  the  respiratory  center  is  also 
increased  by  the  stimulation  of  the  skin  by  the  air,  the  contact  with  clothing, 
and  by  the  hands  of  the  nurse.  We  have  already  seen  that  cutaneous 
stimulation  leads  to  an  increase  in  both  respiratory  rate  and  amplitude  even 
in  the  adult,  a  reaction  that  is  more  pronounced  in  the  child.  The  primary 
stimulus  for  the  establishment  of  the  respiratory  rhythm  at  first,  then,  is  the 
venosity  of  the  blood,  but  this  cause  is  supported  by  the  afferent  cutaneous 
impulses  producing  reflexes  through  the  respiratory  center. 

Certain  Special  Types  of  Respiration. — Whatever  the  exact  quality 
of  the  venous  blood  which  excites  the  respiratory  center  to  produce  normal 
respirations,  there  can  be  no  doubt  that,  as  the  blood  becomes  more  and 
more  venous  from  obstruction  to  the  entrance  of  air  into  the  lungs  or  from 
the  blood  not  taking  up  from  the  air  its  usual  supply  of  oxygen,  the  respi- 
ratory center  becomes  either  less  or  more  active  and  excitable.  Conditions 
ensue  which  have  received  the  names  Apnea,  diminished  breathing;  Hyper- 
pnea,  excessive  breathing;  Dyspnea,  difficult  breathing;  and  Asphyxia, 
suffocation. 


316  RESPIRATION 

Apnea. — This  is  a  condition  of  diminished  respiratory  movement.  When 
we  take  several  deep  inspirations  in  rapid  succession  by  voluntary  effort, 
we  find  that  we  can  do  without  breathing  for  a  much  longer  time  than  usual; 
in  other  words,  several  rapid  respirations  seem  to  inhibit  for  a  time  normal 
respiratory  movements.  The  reason  for  this  partial  cessation  of  respira- 
tion, or  apnea,  is  not  that  we  overcharge  our  blood  with  oxygen,  as  was  once 
thought,  for  Hering  has  shown  that  animals  in  a  condition  of  apnea  may 
have  less  oxygen  in  their  blood  than  in  a  normal  state,  although  the  carbon 
dioxide  is  less.  It  is  probable  that  the  cause  of  apnea  is  that  by  rapid  in- 
flations of  the  lungs  impulses  pass  up  by  the  vagi  by  means  of  which  in- 
spiration is  after  a  while  inhibited;  or  that  by  the  repeated  stimulation  of 
the  center  by  vagus  impulses  which  result  in  rapid  respiratory  movements, 
anabolism  is  at  last  arrested.  Apnea  is  with  difficulty  produced,  if  at  all, 
when  the  vagi  are  divided. 

Asphyxia. — The  condition  of  stress  in  the  respiratory  apparatus  brought 
about  by  insufficient  respiratory  activity  leads  to  a  condition  of  asphyxia. 
Progressive  asphyxiation  may  be  brought  on  by  anything  which  interferes 
with  the  normal  interchange  of  the  respiratory  gases  of  the  blood. 

Asphyxia  may  be  produced  by  the  prevention  of  the  due  entry  of  oxygen 
into  the  blood,  either  by  direct  obstruction  of  the  trachea  or  other  part  of  the 
respiratory  passages,  or  by  introducing  instead  of  ordinary  air  a  gas  devoid 
of  oxygen,  or  by  interference  with  the  due  interchange  of  gases  between 
the  air  and  the  blood. 

The  respiratory  symptoms  of  progressive  asphyxiation  may  be  divided 
into  three  groups,  which  correspond  with  the  stages  of  the  condition  most 
readily  recognized;  these  are:  i,  the  stage  of  exaggerated  breathing,  hyperpnea; 
2,  the  stage  of  convulsions,  dyspnea;  3,  the  stage  of  exhaustion,  asphyxiation. 

In  the  first  stage  the  breathing  becomes  more  rapid  and  at  the  same  time 
deeper  than  usual,  the  inspirations  at  first  being  especially  exaggerated  and 
prolonged.  This  is  soon  followed  by  a  similar  increase  in  the  expiratory 
efforts  being  aided  by  the  muscles  of  extraordinary  expiration.  This  stage  is 
usually  called  hyperpnea.  Hyperpnea  soon  passes  into  a  condition  of  labored 
breathing  in  which  there  is  marked  increase  of  the  force  of  the  expiratory 
as  well  as  of  the  inspiratory  act,  a  condition  described  as  dyspnea.  All  the 
muscles  capable  of  aiding  either  directly  or  indirectly  in  respiration 
are  ultimately  brought  into  action.  These  respiratory  convulsions  are 
followed  by  rather  sudden  onset  of  paralysis  of  the  respiratory  center  and 
ultimate  death. 

The  conditions  of  the  vascular  system  in  asphyxia  are:  i,  more  or  less 
interference  with  the  passage  of  the  blood  through  the  systemic  and  the  pul- 
monary blood  vessels;  2,  accumulation  of  blood  in  the  right  side  of  the  heart 
and  in  the  systemic  veins;  3,  circulation  of  impure  (non-aerated)  blood  in 
all  parts  of  the  body,  especially  through  the  respiratory  center;  4,  great 


EFFECTS    OF  VITIATED    AIR  317 

slowing  of  the  heart  by  stimulation  of  the  vagus  center  from  lack  of 
oxygen. 

It  must  be  kept  clearly  in  mind  that  the  respiratory  changes  just 
described  as  characteristic  of  asphyxiation  are  the  secondary  results  of  the 
primary  general  tissue  asphyxiation.  If  an  animal  is  deprived  of  its  income 
of  oxygen,  and  its  ability  to  eliminate  the  product  of  oxidation,  carbon 
dioxide,  the  nutritive  balance  around  the  peripheral  tissues  is  immediately 
disturbed.  The  result  is  that  tissue  metabolism  such  as  occurs  is  deranged. 
If  oxidations  are  incomplete,  intermediary  products  accumulate  and  on 
the  whole  the  physiological  life  of  the  tissue  is  rapidly  blocked.  The 
average  organ  of  the  human  body  can  endure  only  a  certain  degree  of  as- 
phyxiation before  changes  occur  which  destroy,  in  whole  or  in  part,  the 
protoplasmic  organization.  Of  all  the  tissues  the  nervous  tissues  are  most 
susceptible  to  asphyxiation.  The  generalized  tissues,  the  epidermis,  con- 
nective tissue,  etc.,  are  most  resistant.  Life  is  jeopardized  by  injury  to  the 
weakest  point,  hence  the  body  as  a  whole  will  not  recover  from  complete 
asphyxiation  which  endures  for  a  time  greater  than  that  which  the  nervous 
tissues  will  withstand.  Stewart,  Guthrie,  Burns  and  Pike  have  set  the 
limits  very  low  for  this  tissue,  from  7  to  16.5  minutes.  However,  incomplete 
asphyxiation  is  a  condition  difficult  to  determine,  and  the  less  complete  the 
asphyxiation,  the  greater  the  probability  of  recovering  the  normal  tissue 
activity.  In  all  those  conditions  of  life  in  which  accidental  asphyxiation 
occurs,  it  must  be  assumed  that  we  are  dealing  with  one  of  partial  asphyxia- 
tion, especially  in  all  efforts  at  resuscitation. 

Cheyne-Stokes'  breathing  is  a  rhythmical  irregularity  in  respirations  which 
has  been  observed  in  various  diseases.  Respirations  occur  in  groups.  At 
the  beginning  of  each  group  the  inspirations  are  very  shallow,  but  each 
successive  breath  is  deeper  than  the  preceding,  until  a  climax  is  reached. 
The  inspirations  then  become  less  and  less  deep,  until  they  cease  altogether 
for  a  time,  after  which  the  cycle  is  repeated.  This  phenomenon  appears  to  be 
due  to  the  want  of  action  of  some  of  the  usual  cerebral  influences  which  pass 
to,  and  regulate  the  discharges  of,  the  respiratory  center. 

Effects  of  Vitiated  Air. — Ventilation. — As  the  air  expired  from  the 
lungs  contains  a  large  proportion  of  carbon  dioxide  and  a  minute  amount 
of  organic  matter,  it  is  obvious  that  if  the  same  air  be  breathed  again  and 
again,  the  proportion  of  carbon  dioxide  and  organic  matter  in  it  will  con- 
stantly increase  till  it  becomes  unfit  to  breathe;  long  before  this  point  is 
reached,  however,  sensations  of  uneasiness  occur,  such  as  headache,  languor, 
and  a  sense  of  oppression.  It  is  a  remarkable  fact,  however,  that  the  organ- 
ism after  a  time  adapts  itself  to  a  very  vitiated  atmosphere,  and  that  a  person 
soon  comes  to  breathe,  without  sensible  inconvenience,  an  atmosphere  which, 
when  he  first  enters  it,  feels  intolerable.  Such  an  adaptation,  however,  can 
take  place  only  at  the  expense  of  a  depression  of  all  the  vital  functions,  which 


318  RESPIRATION 

must  be  injurious  if  long-continued  or  often  repeated.  This  power  of  adapta- 
tion is  well  illustrated  by  an  experiment  of  Claude  Bernard.  If  a  sparrow 
is  placed  under  a  bell-glass  of  such  size  that  it  will  live  for  three  hours,  be 
taken  out  at  the  end  of  the  second  hour  (when  it  could  have  survived  another 
hour),  and  a  fresh  healthy  sparrow  introduced,  the  latter  will  die  at  once. 

It  must  be  evident  that  provision  for  a  constant  and  plentiful  supply  of 
fresh  air,  and  the  removal  of  that  which  is  vitiated,  are  of  greater  importance 
than  the  actual  cubic  space  per  person  of  occupants.  Not  less  than  2,000 
cubic  feet  per  individual  should  be  allowed  in  sleeping  apartments  (bar- 
racks, hospitals,  etc.),  and  with  this  allowance  the  air  can  be  maintained  at 
the  proper  standard  of  purity  only  by  such  a  system  of  ventilation  as  pro- 
vides for  the  supply  of  1,500  to  2,000  cubic  feet  of  fresh  air  per  person  per 
hour. 

Efects  of  Breathing  Gases  Other  than  the  Atmosphere. — Asphyxiation  is 
produced  by  the  direct  poisonous  action  of  such  gases  as  carbon  monoxide, 
which  is  contained  to  a  considerable  amount  in  common  coal  gas.  The 
fatal  effects  often  produced  by  this  gas  (as  accidents  from  burning  charcoal 
stoves  in  small,  close  rooms)  are  due  to  its  entering  into  combinations  with 
the  hemoglobin  of  the  blood  corpuscles  and  thus  preventing  the  formation 
of  oxyhemoglobin  because  of  the  more  stable  carbon- monoxide  hemoglobin. 
The  partial  pressure  of  oxygen  in  the  atmosphere  may  be  considerably  in- 
creased without  much  effect  in  displacing  the  carbon  monoxide,  hence  this 
is  rapidly  fatal  when  breathed.  Hydrogen  may  take  the  place  of  nitrogen 
with  no  marked  ill  effect,  if  the  oxygen  is  in  the  usual  proportions.  Sul- 
phureted  hydrogen  destroys  the  hemoglobin  of  blood  and  thus  produces  oxygen 
starvation.  Nitrous  oxide  acts  directly  on  the  nervous  system  as  a  narcotic, 
and  may  also  form  a  compound  with  hemoglobin.  Certain  gases,  such  as 
carbon  dioxide  in  more  than  a  certain  proportion,  sulphurous  acid  gases,  am- 
monia, and  chlorine,  produce  spasmodic  closure  of  the  glottis  and  prevent 
respiration. 

Alteration  in  the  Atmospheric  Pressure. — Lower  barometric  pres- 
sures than  the  normal  occur  in  high  altitudes,  for  example  in  mountain 
climbing  or  in  aerial  navigation.  The  susceptibility  to  decrease  in  baro- 
metric pressure  varies  in  different  individuals.  At  an  altitude  of  about 
10,000  feet  many  persons  begin  to  experience  mountain  sickness,  though 
most  individuals  are  not  so  affected  until  they  ascend  to  15,000  feet  or 
more.  The  symptoms  that  develop  are  nausea,  dizziness,  palpitation  of 
the  heart,  headache,  and  muscular  weakness.  The  oxygen  partial  pres- 
sure of  the  atmosphere  is  reduced  to  half  at  about  15,000  feet  elevation. 
At  this  pressure  the  body  begins  to  show  some  stress  from  inability  to 
get  an  adequate  quantity  of  oxygen.  The  tension  of  the  oxygen  in  the 
alveolar  air  is  not  great  enough,  see  figure  238  showing  the  relation  of  the 
partial  pressure  of  oxygen  and  the  percentage  of  hemoglobin  saturation, 


ALTERATION    IN    THE    ATMOSPHERIC    PRESSURE 


to  allow  the  blood  of  the  pulmonary  capillaries  to  combine  with  its  usual 
quantity  of  oxygen.  There  is  enough  oxygen  absorbed,  however,  to 
satisfy  the  amount  used  by  the  tissues  under  ordinary  circumstances. 
It  is  only  when  an  extra  amount  of  activity  is  called  for  that  stress  is 
observed  at  this  level.  At  still  greater  altitudes  the  oxygen  of  the  arterial 
blood  is  further  reduced  until  a  level  is  reached  at  which  the  total  amount 
of  oxygen  absorbed  by  the  pulmonary  blood  is  less  than  that  normally 
lost  in  the  tissues.  This  produces  a  real  tissue  oxygen  want.  The  con- 
condition  receives  the  technical  name  anoxemia. 

Progressive  anoxemia  sets  into  activity  a  number  of  physiological 
mechanisms  which  aid  the  body  to  absorb  its  maximum  of  oxygen  from  the 
alveolar  air.  These  reactions  are  called  compensatory.  They  have  been 
described  by  a  number  of  workers  in  the  Medical  Division  of  the  United 
States  Army  who  developed  a  technique  for  testing  the  ability  of  the 


FIG.  2410. — The  progressive  increase  of  the  amount  of  hemoglobin  in  the  blood 
during  a  journey  from  England  to  the  high  Andes  (Richards). 

human  body  to  withstand  low  atmospheric  pressures  in  aviation.  The 
compensatory  factors  lead  to  great  increase  in  respiratory  rate,  an  increase 
in  the  tidal  air,  therefore,  a  great  increase  in  the  respiratory  minute 
volume  of  air  breathed.  The  heart  rate  is  also  greatly  increased  thus 
maintaining  a  higher  systolic  blood  pressure  notwithstanding  the  fact 
that  there  is  vascular  dilation,  therefore,  increased  volume  of  blood  flow. 
Also,  the  percentage  of  hemoglobin  in  the  blood  is  increased  if  the  low 
oxygen  pressure  acts  through  sufficient  time,  as  in  mountain  residence,  figure 
2410.  When  the  limit  of  compensation  is  reached,  then  the  body  quickly 
succumbs  through  the  following  symptoms.  Respiration  decreases  in  rate 
and  in  amplitude  and  stops.  The  blood  pressure  at  first  becomes  high, 
then  falls  slowly  at  first,  but  more  rapidly  later  to  a  final  low  level.  The 
heart  rate,  which  is  greatly  accelerated  in  the  early  stage  of  anoxemia,  is 
enormously  slowed  in  the  late  stages  and  especially  at  the  time  of  and 
following  the  stopping  of  respiration.  It  would  seem  that  the  lack  of 
oxygen  at  first  strongly  stimulates  the  medullary  centers  of  respiration,  of 
cardiac  acceleration,  and  of  vaso  constriction.  But  in  extreme  anoxemia 


320 


RESPIRATION 


the  respiratory  center  is  no  longer  supported  in  activity,  and  the  cardiac 
inhibitory  center  is  stimulated  to  inhibitory  spasm.  This  is  shown  in 
figure  2416.  This  figure  represents  only  the  terminal  effect  of  anoxemia 
in  the  dog  under  chloretone  anesthesia.  The  stopping  of  respiration  is 
shown  in  the  top  trace.  The  blood  pressure  tracing  shows  an  enormous 
slowing  of  the  heart  rate  by  anoxemial  stimulation  of  the  vagus  medullary 
center.  The  proof  is  found  in  the  rapid  heart  rate  after  the  vagus  nerves 
are  both  cut. 


' 


FIG.  2416. — The  effects  of  extreme  anoxemia  in  the  dog. 

The  compensatory  increase  in  respiratory  rate  and  volume  during 
oxygen  want  disturbs  the  factor  of  carbon  dioxide  balance  in  the  blood. 
The  carbon  dioxide  is  lost  more  rapidly  than  usual,  hence  its  concentration 
in  the  blood  is  diminished,  often  from  twenty  to  fifty  per  cent.  That 
carbon  dioxide  is  the  chemical  stimulator  of  the  respiratory  center  has 
long  been  known,  but  Henderson  has  more  recently  given  evidence  of  the 
stimulating  effect  of  carbon  dioxide  on  certain  other  functions  of  the 
nervous  and  muscular  mechanisms.  He  has  called  the  condition  of  reduced 
carbon  dioxide  acapnia,  and  offers  the  suggestion  that  lack  of  sufficient 
carbon  dioxide  may  contribute  to  the  complex  of  symptoms  associated 
with  coincident  lack  of  oxygen. 

Men  are  often  subjected  to  higher  than  normal  barometric  pressures  in 
caisson  work,  diving,  etc.  Paul  Bert  has  found  in  experimenting  with 
animals  that  the  oxygen  pressures  may  be  gradually  increased  to  a  con- 
siderable extent  without  marked  effect,  even  to  the  extent  of  8  or  10  atmos- 
pheres, but  when  the  oxygen  pressure  is  increased  up  to  20  atmospheres  the 
oxygen  becomes  poisonous  and  the  animals  experimented  upon  died  with 


RESUSCITATION    FROM    ELECTRIC    SHOCK    AND    DROWNING       321 

severe  tetanic  convulsions.  However,  caisson  workers  often  experience 
very  severe  symptoms,  such  as  bleeding  from  the  nose,  dyspnea,  vascular 
inco-ordination,  etc.  These  symptoms  are  due  not  so  much  to  the  great 
increase  in  pressure  as  to  the  release  from  the  pressure.  When  the 
pressure  is  released  too  rapidly,  the  excess  of  gases  in  the  tissues  and  in  the 
blood  are  set  free  more  rapidly  than  they  can  be  thrown  off  by  excretion 
processes.  Gases,  as  such,  gather  in  the  blood  vessels  and  form  embolisms 
which  occlude  the  finer  vessels.  This,  of  course,  produces  serious  dis- 
turbances in  the  nutrition  of  the  parts  involved.  If  these  parts  happen  to 
be  vital,  death  may  result. 

Resuscitation  from  Electric  Shock  and  Drowning. — Of  the  numerous 
conditions  which  lead  to  accidental  asphyxiation,  electric  shock  on  the  one 
hand,  and  drowning  on  the  other  are  of  great  scientific  and  practical  importance 
in  the  present  day.  These  special  conditions  of  asphyxiation  necessarily  in- 
volve problems  of  general  tissue  asphyxiation  and  resuscitation.  Under  the 
influence  of  electric  shock  of  sufficient  intensity,  an  immediate  result  is 
paralysis  of  the  nervous  respiratory  control,  with  whatever  else  may  be 
directly  or  indirectly  involved.  This  condition  quickly  brings  on  asphyxia- 
tion with  all  of  its  train  of  perverted  functional  activity.  So  also  in  drowning, 
suspension  under  water  blocks  respiratory  activity  and  induces  asphyxia- 
tion. Within  the  last  few  years  careful  investigation  of  this  condition  has 
been  made  by  Stewart,  Guthrie  and  Pike,  by  Crile  and  Dolley,  and  by 
numerous  others.  The  work  has  tended  to  set  the  time  limits  of  tissue 
asphyxiation  after  which  recovery  is  impossible  or  at  most  incomplete.  The 
nerve  tissues  are  most  susceptible  to  injury  here,  see  page  303.  Within 
the  nervous  tissues,  the  different  functional  centers  manifest  different  degrees 
of  susceptibility.  Those  immediately  involved  in  the  injuries  are  the  re- 
spiratory, vaso-motor,  and  cardiac  regulative  centers.  If  all  the  activities 
of  these  nervous  mechanisms  can  be  re-established,  control  of  general  vis- 
ceral reactions  will  be  insured.  The  delicate  functional  activity  of  the 
higher  or  cortical  regions  of  the  brain  are  even  more  susceptible  to  asphyxia- 
tion, and  recover  more  slowly  if  at  all. 

In  the  condition  of  drowning  there  are  no  highly  important  special 
injuries.  In  electric  shock  on  the  other  hand,  there  may  be  a  series  of 
local  injuries  from  electric  burns,  etc.  These  may  injure  only  the  point  of 
contact  between  the  surface  of  the  body  and  the  electric  conductor,  but  it 
is  perfectly  possible  that  the  injury  may  be  intense  on  some  deep-seated 
vital  structure.  In  such  cases,  recovery  of  the  respiration  or  of  the  circula- 
tion will  not  necessarily  insure  ultimate  success  in  the  efforts  to  revive  the 
individual. 

Considering  the  fact  that  the  nervous  tissue  cannot  be  safely  recovered 
beyond  the  limit  of  15  minutes  (this  is  a  fair  maximum  average  from  our 
most  reliable  authorities)  it  follows  that  immediate  and  careful  steps  must 


322  RESPIRATION 

be  taken  to  eliminate  the  conditions  producing  tissue  asphyxiation,  i.e., 
to  re-establish  both  respiration  and  circulation.  Artificial  respiration  in  one 
form  or  another  is  the  first  aid  to  be  given  in  drowning,  and  in  other  types  of 
asphyxiation,  since  the  technique  is  equally  effective  after  the  lungs  are 
emptied  of  fluid.  The  method  at  present  most  relied  upon  is  that  of  Schafer 
which  includes  both  the  artificial  respiration  and  indirect  heart  massage. 
The  procedure  to  be  followed  in  Schafer' s  method  in  condensed  statement 
is  given  as  follows  by  Dolley: 

"  The  patient  is  rolled  upon  his  belly,  the  face  turned  to  one  side,  and  the 
arms  are  extended  as  straight  forward  as  possible.  The  extension  of  the 
arms  is  a  very  important  improvement,  introduced  by  the  Commission,  on 
the  original  Schafer  method.  The  operator  kneels  straddling  the  patient's 
thighs  and  facing  his  head;  he  places  his  palms  on  the  muscles  of  the  small 
of  the  back  with  the  fingers  spread  over  the  lowest  ribs.  Then  holding  his 
arms  straight,  he  swings  forward  so  that  his  weight  is  gradually  brought  to 
bear  upon  the  subject.  This  should  take  from  two  to  three  seconds,  and  must 
not  be  violent.  It  compresses  both  the  chest  and  the  abdomen.  The  result 
is  that  not  only  is  the  chest  compressed  from  front  to  back,  but  the  pressure 
on  the  abdominal  viscera  tends  to  force  the  diaphragm  upward.  The  air  is 
forced  out  of  the  lungs,  expiration.  The  operator  then  immediately  swings 
back  to  his  starting  position.  Through  their  elasticity  the  chest  walls  ex- 
pand and  air  is  inspired.  A  two-second  interval  should  follow  the  forced 
expiration  so  that  the  rate  is  from  twelve  to  fifteen  a  minute.  The  method 
not  only  accomplishes  safely  and  easily  ventilation  of  the  lungs,  but  it  must 
affect  a  fair  amount  of  compression  and  relaxation  of  the  heart,  especially 
in  young  or  thin  individuals.  This  so-called  indirect  heart  massage,  which 
will  be  more  emphasized  later,  is  a  valuable  stimulant  to  a  failing  heart." 
Artificial  respiration  should  be  kept  up  for  from  two  to  four  hours.  A 
slight  but  temporary  circulation  of  the  blood  may  produce  a  partial  oxidation 
which  only  very  slowly  recovers  sufficient  vital  activity  to  bring  the  nerve 
centers  up  to  the  automatic  and  reflex  level  of  activity  required. 

Some  cases  of  so-called  drowning  are  in  reality  death  from  cardiac  failure. 
Any  hope  of  resuscitation  in  this  type  depends  upon  vigorous  indirect 
cardiac  massage.  This  is  accomplished  more  effectively  in  the  above  method 
of  artificial  respiration  by  allowing  the  palms  of  the  hands  to  slide  around 
to  the  sides  of  the  body,  presssing  near  the  ends  of  the  free  ribs.  In  this 
position  it  is  easy  to  give  pressure  with  the  finger  tips  under  the  ribs  and 
against  the  heart.  In  extreme,  and  perhaps  in  surgical  cases,  direct  massage 
may  be  given.1 

1  Fuller  discussion  of  the  conditions  involving  drowning  and  the  procedure  look- 
ing toward  recovery  are  available  in  the  following  references:  C.  C.  Guthrie,  Blood- 
vessel Surgery,  chapter  on  Resuscitation,  page  300.  Report  of  the  Commission  on 
Resuscitation  from  Electric  Shock,  W.  B.  Cannon,  Chairman.  Medical  Handbook 
for  the  Use  of  Lighthouse  Vessels,  etc.,  published  by  the  U.  S.  Public  Health  and 
Marine-Hospital  Service.  D.  H.  Dolley,  On  Resuscitation,  Bulletin  of  the  University 
of  Missouri,  Medical  Series,  No.  4. 


EFFECT    OF    RESPIRATION    ON    THE    CIRCULATION 


323 


Apparent  recuperation  with  the  re-establishment  of  both  normal  and 
respiratory  activity  may  occur,  yet  later  stoppage  will  come  about.  Prolonged 
asphyxiation  leaves  the  body  and  tissues  so  clogged  with  carbon  dioxide 
and  other  waste  products,  that  the  renewal  of  the  vital  activity  of  the  nervous 
centers  is  under  a  weakened  condition.  In  such  cases  it  is  highly  necessary 
to  prolong  artificial  respiration.  Even  after  asphyxiation  of  short  duration 
it  may  be  some  hours  or  even  days  before  the  body  is  brought  back  to  its 
normal  level  of  functional  efficiency. 

THE  EFFECT  OF  RESPIRATION  ON  THE  CIRCULATION. 

As  the  heart,  the  aorta,  and  pulmonary  vessels  are  situated  in  the  air- 
tight thorax,  they  are  exposed  to  a  certain  alteration  of  pressure  when  the 
capacity  of  the  latter  is  varied  during  respiration.  The  disturbance  of  pres- 
sure which  occurs  during  inspiration  causes,  first,  a  .decrease  in  the  intra- 


FIG.  242. — Diagram  of  an  Apparatus  Illustrating  the  Effect  of  Inspiration  upon  the 
Heart  and  Great  Vessels  within  the  Thorax.  I,  The  thorax  at  rest;  II,  during  inspiration; 
D  represents  the  diaphragm  when  relaxed;  D',  when  contracted  (it  must  be  remembered 
that  this  position  is  a  mere  diagram),  i.  e.,  when  the  capacity  of  the  thorax  is  enlarged;  H, 
the  heart;  V,  the  veins  entering  it,  and  A,  the  aorta;  Rl,Ll,  the  right  and  left  lung;  T,  the 
trachea;  M,  mercurial  manometer  in  connection  with  pleura.  The  increase  in  the  capacity 
of  the  box  representing  the  thorax  is  seen  to  dilate  the  heart  as  well  as  the  lungs,  and  so  to 
pump  in  blood  through  V,  whereas  the  valve  prevents  reflux  through  A.  The  position  of 
the  mercury  in  M  shows  also  the  suction  which  is  taking  place.  (Landois.) 


thoracic  pressure,  a  decrease  which  affects  all  the  organs  of  the  thorax — the 
lungs,  the  great  blood-vessels,  the  heart.  The  expansion  of  the  elastic  lungs 
counterbalances  this  change  in  pressure  in  part,  but  it  never  does  so  entirely, 
since  part  of  the  pressure  within  the  lungs  is  expended  in  overcoming  their 
elasticity.  The  amount  thus  used  up  increases  as  the  lungs  become  more 
and  more  stretched,  so  that  the  intrathoracic  pressure  during  inspiration,  as 


324  RESPIRATION 

far  as  the  heart  and  great  vessels  are  concerned,  never  quite  equals  the  intra- 
pulmonary  pressure,  and  at  the  conclusion  of  inspiration  is  considerably 
less  than  the  atmospheric  pressure.  It  has  been  ascertained  that  the  amount 
of  the  pressure  used  up  in  the  way  above  described  varies  from  5  to  7  mm.  of 
mercury  in  ordinary  inspiration,  to  30  mm.  of  mercury  at  the  end  of  a  deep 
inspiration.  So  it  will  be  understood  that  the  pressure  to  which  the  heart 
and  great  vessels  are  subjected  diminishes  as  inspiration  progresses,  and  at 
its  summit  is  less  by  from  7  to  30  mm.  than  the  normal  atmospheric  pres- 
sure, 760  mm.  of  mercury.  It  will  be  understood  from  the  accompanying 
diagram  how  an  increase  in  the  volume  of  the  thorax  will  have  the  effect  of 
pumping  blood  into  the  heart  from  the  veins.  During  inspiration  the  pres- 
sure outside  the  heart  and  great  vessels  is  diminished,  and  they,  by  virtue  of 
their  elasticity,  have  therefore  a  tendency  to  expand  and  to  diminish  the  intra- 
vascular  pressure.  The  diminution  of  pressure  within  the  veins  passing 
to  the  right  auricle  and  within  the  right  auricle  itself,  will  draw  the  blood 
into  the  thorax,  and  so  assist  the  circulation.  This  suction  action  of  the 
thorax  is  the  cause  of  the  slight  negative  pressure  of  the  ventricles  previously 
described.  The  effect  of  more  blood  in  the  right  auricle  will,  cateris  paribus, 
increase  the  amount  passing  through  the  right  ventricle,  and  through  the 
lungs  into  the  left  auricle  and  ventricle,  and  thus  into  the  aorta.  This  all 
tends  to  increase  the  blood-pressure.  The  effect  of  the  diminished  pressure 
upon  the  pulmonary  vessels  will  also  help  toward  the  same  end,  an  increased 
flow  through  the  lungs,  so  that,  as  far  as  the  mechanical  effects  on  the  heart 
and  its  veins  are  concerned,  inspiration  increases  the  blood-pressure  in  the 
arteries.  The  effect  of  inspiration  upon  the  aorta  and  its  branches  within 
the  thorax  would  be,  however,  contrary;  for  as  the  external  pressure  is  dimin- 
ished, the  vessels  would  tend  to  expand,  and  thus  to  diminish  the  tension  of 
the  blood  within  them,  but,  inasmuch  as  the  relative  variation  in  pressure 
on  the  large  arteries  is  slight,  the  diminution  of  arterial  tension  caused  by 
this  means  will  be  insufficient  to  counteract  the  increase  of  blood-pressure 
produced  by  the  effect  of  inspiration  upon  the  volume  of  discharge  of  the 
veins  of  the  chest,  and  the  balance  of  the  whole  action  would  be  in  favor  of 
an  increase  of  blood-pressure  during  the  inspiratory  period.  When  a  blood- 
pressure  tracing  is  taken  at  the  same  time  that  the  respiratory  movements 
are  being  recorded,  it  will  be  found  that,  although,  speaking  generally,  the 
arterial  tension  is  increased  during  inspiration,  the  maximum  of  arterial 
tension  does  not  correspond  with  the  acme  of  inspiration,  figure  243.  In 
fact,  at  the  beginning  of  inspiration  the  pressure  continues  to  fall  for  a  brief 
moment,  then  gradually  rises  until  the  end  of  inspiration,  and  continues  to 
do  so  for  a  moment  after  expiration  has  commenced.  For  explanation  of 
the  influence  of  heart  rate  in  this  variation  of  blood-pressure,  associated 
with  the  respiratory  movement,  see  page  212. 

In  ordinary  expiration  all  this  would  be  reversed,  but  if  the  abdominal 


EFFECT    OF    RESPIRATION    ON    THE    CIRCULATION  325 

muscles  are  violently  contracted,  as  in  extraordinary  expiration,  the  same 
relative  effect  would  be  produced  as  by  inspiration.  The  immediate  effect 
during  inspiration  of  the  diminished  intra-thoracic  pressure  upon  the  pul- 
monary vessels  is  to  produce  an  initial  dilatation  of  both  artery  and  veins, 
and  this  delays  for  a  moment  the  passage  of  blood  toward  the  left  side  of 
the  heart,  resulting  in  an  initial  fall  in  the  arterial  pressure,  but  the  fall  of 
blood-pressure  is  immediately  followed  by  a  steady  rise,  since  the  flow  is 
increased  by  the  initial  dilatation  of  the  vessels.  The  converse  is  the  case 
with  expiration.  As,  however,  the  pulmonary  veins  are  more  easily  di- 
latable than  the  pulmonary  artery,  their  greater  distensibility  increases  the 


FIG.  243. — Comparison  of  Blood-pressure  Curve  with  Curve  of  Intra-thoracic  Pressure. 
(To  be  read  from  left  to  right.)  a  is  the  curve  of  blood-pressure  with  its  respiratory 
undulations,  the  slower  beats  on  the  descent  being  very  marked;  b  is  the  curve  of  intra- 
thoracic  pressure  obtained  by  connecting  one  limb  of  a  manometer  with  the  pleural  cavity. 
Inspiration  begins  at  i  and  expiration  at  e.  The  intra-thoracic  pressure  rises  very  rapidly 
after  the  cessation  of  the  inspiratory  effort,  and  then  slowly  falls  as  the  air  issues  from 
the  chest;  at  the  beginning  of  the  inspiratory  effort  the  fall  becomes  more  rapid.  (M. 
Foster.) 

flow  of  blood  as  inspiration  proceeds,  while  during  expiration,  except  at  its 
beginning,  this  property  of  theirs  acts  in  the  opposite  direction,  and  diminishes 
the  flow.  Thus,  at  the  beginning  of  inspiration  the  diminution  of  blood 
pressure,  which  commenced  during  expiration,  is  continued,  but  after  a  time 
the  diminution  is  succeeded  by  a  steady  rise.  The  reverse  is  the  case  with 
expiration,  i.e.,  there  is  at  first  a  rise  and  then  a  fall  of  blood  pressure. 

As  regards  the  effect  of  expiration,  the  capacity  of  the  chest  is  diminished 
and  the  intra-thoracic  pressure  returns  to  the  normal,  which  is  still  slightly 
below  the  atmospheric  pressure.  The  effect  of  this  on  the  veins  is  to  in- 
crease their  extravascular  and  so  their  intravascular  pressure,  and  to  di- 
minish the  flow  of  blood  into  the  left  side  of  the  heart.  This  will,  of  course, 
react  to  decrease  the  general  blood-pressure.  Ordinary  expiration  does  not 
produce  a  distinct  obstruction  to  the  circulation,  as  even  when  the  expiration 
is  at  an  end  the  intra-thoracic  pressure  is  less  than  the  atmospheric  pressure. 
The  effect  of  violent  expiratory  efforts,  however,  does  have  a  distinct  action 


326  RESPIRATION 

in  obstructing  the  current  of  blood  through  the  lungs,  as  seen  in  the  con- 
gestion in  the  exaggerated  condition  of  straining,  this  condition  being  pro- 
duced by  pressure  on  the  entire  group  of  pulmonary  vessels. 

There  are  other  mechanical  factors,  such,  for  example,  as  the  effect  of 
the  abdominal  movements,  both  in  inspiration  and  in  expiration,  upon  the 
arteries  and  veins  within  the  abdomen  and  of  the  lower  extremities.  Also 
the  influence  of  the  varying  intrathoracic  pressure  upon  the  pulmonary 
vessels,  which  ought  to  be  taken  into  consideration.  The  effect  of  the 
abdominal  movements  during  inspiration  is  twofold.  On  the  one  hand, 
blood  is  sent  upward  into  the  chest  by  compression  of  the  vena  cava  inferior; 
on  the  other  hand,  the  passage  of  blood  downward  from  the  chest  through 
the  abdominal  aorta,  and  upward  in  the  veins  of  the  lower  extremity,  is  to 
a  certain  extent  obstructed. 


327 


LABORATORY  EXPERIMENTS  IN  RESPIRATION. 

1.  Respiratory  Rate  in  Man.  —  Count  your  respirations  for  from  2 
to  4  minutes  while  sitting  quietly,  and  determine  the  average  number  per 
minute.     Repeat  the  counting  after  standing  for  5  minutes,  and  after 
brisk  exercise.     These  determinations  involve  the  element  of  conscious- 
ness, under  which  condition  it  is  difficult  for  a  person  to  breathe  with  his 
normal  rate  and  depth. 

Make  a  series  of  determinations  of  respiratory 
rates  of  persons  who  are  sitting  quietly  but  uncon- 
scious of  your  determinations.  Count  the  rates  in 
a  number  of  persons  of  different  ages;  where  possible, 
take  into  consideration  height,  weight,  etc.  Tabu- 
late the  results  for  a  comparison  and  for  future 
reference. 

2.  The  Character  of  Respiratory  Movements 
in   Man.  —  A   number  of  instruments  have  been 
devised  for  measuring  human   respiratory  move- 
ment,   many    of    which    measure    the    change    in 
diameter   of   the  chest  in  respiratory    movement. 
Adjust  one  of  these,  for  example  Burdon-Sander- 
son's  stethograph,  to  the  thorax,  and  record  the 
movement  of  the  receiving  tambour  on  a  smoked- 

paper  kymograph  which  travels  at  the  rate  of  i 

,       m,  .  ,    j  FIG.  244.  —  Change  in 

cm.  per  second.     Inis  record,  called  a  stethogram,   Diameter  of  the  Body  in 

will  exhibit  the  respiratory  rate,  the  relative  time  of  Respiration,  Costal  Type. 
^,       >.  ,  ,  ,    ^,       a.  Outline  of  the  body  in 

the  linspiratory   and   expiratory  phases,   and  the  forced  expiration.    In  the 

character   of  each.  heavy  continuous  line,  b, 


3.  The  Actual  Change  of  Diameter  in  the  Chest 
in  Respiration.  —  Use  a  caliper  provided  for  the  pur-  ordinary  inspiration  and 

,       i  ,    ,.  f    ,       the  inner  margin  that  of 

pose  and  measure  the  dorso-  ventral  diameter  of  the  ordinary  expiration.     ct 

chest  at  a  series  of  points  along  the  sternum,  taking  Contour  of  forced  inspira- 
j.  J_1      ,     .   ,  ^      ,.  Al  .  ,  tion.  (After  Hutchinson.) 

the  reading  at  the  height  of  the  inspiratory  phase 

and  of  the  expiratory  phase  in  ordinary  respiration.  Repeat  the  measure- 
ment in  forced  respiration.  Map  the  results  on  millimeter  paper,  as 
indicated  in  figure  244. 

Repeat  these  measurements  in  the  transverse  diameter  at  the  first, 
fifth,  and  tenth  ribs. 

Using  the  thoracograph,  figure  245,  record  the  outline  of  the  cross 
section  of  the  chest  at  the  level  of  the  mammae,  tenth  rib,  and  the  umbili- 
cus, showing  the  volume  changes  in  the  following  four  positions:  (i) 
Ordinary  expiration,  (2)  ordinary  inspiration,  (3)  forced  expiration,  (4) 
forced  inspiration,  see  figure  246. 


RESPIRATION 


4.  The  Volume  of  Air  Breathed  by  Man. — Determine  the  average  vol- 
ume of  air  breathed  per  respiration,  using  Hutchinson's  spirometer,  figure 
235,  set  the  instrument  at  the  zero  point,  exhale  into  the  instrument 

through  the  tube,  using  all  possible  care 
Ml   to  breathe  with  your  normal  rate  and 

I 

;'  ~      ;        depth.     Better  results  will  be  obtained 

dgfe|&j  by  taking  the  average  from  sets  of  ten 

j       ?  consecutive  expirations  into  the  instru- 

^^  ment.     From  the  average  of  the  volume 

per  respiration,  and  the  average  number 
of  respirations  per  minute,  determined 
in  Experiment  i,  calculate  the  amount 
of  air  breathed  per  hour  and  per  day. 
5 .  Vital  Capacities. — Using  the  spiro- 
meter as  in  the  preceding  experiment, 
set  the  instrument  at  zero  and  exhale 
into  it: 

a.  Begin   with   the   fullest   possible 
inspiration  and  exhale  the  greatest  pos- 
sible amount  of  air  from  the  lungs.     This 
is  known  as  the  vital  capacity. 

b.  Beginning  at  the  end  of  an  ordi- 
FIG.  245.— Thoracograph  of  Deufestcl.  naiT  expiration  exhale  into  the  instru- 
ment   the    greatest    possible    amount. 

This  is  called  the  reserve  air. 

c.  Following  ordinary  inspiration  exhale  into  the  instrument  until  you 
reach  the  ordinary  state  of  expiration.     This  involves  the  conscious  fixing 
of  two  points  in  the  respiratory  act,  namely,  the  summit  of  inspiration  and 
of  expiration,  which  are  ordinarily  automatically  adjusted  by  the  body. 
The  error  of  the  determination  is  therefore  great.     It  is  better  to  make  this 
measurement  in  sets  of  ten,  as  in  the  preceding  experiment,  and  take  the 
average.     This  reduces  the  error.     This  quantity  of  air  is  known  as  the 
tidal  air.     One  can  measure  the  tidal  air  and  the  reserve  air  together,  check 
them  against  the  sum  of  the  two,  as  in  a  and  b. 

d.  The  sum  of  the  tidal  and  reserve  air  taken  from  the  vital  air  will 
leave  the  amount  which  one  may  inspire  over  and  above  that  in  the  chest 
at  the  end  of  ordinary  inspiration.     This  is  called  complemental.     The 
complemental  can  be  measured  by  inspiring  the  air  from  the  spirometer, 
but  this  is  not  good  hygienic  practice  where  large  numbers  are  using  the 
same  instrument,  unless  the  instrument  be  thoroughly  cleaned  before  the 
inspiration  is  taken. 

6.  The  Respiratory  Pressure  in  Man. — Measure  there  spiratory 
pressure,  the  variation  in  pressure  of  the  air  in  the  air-passages,  by  means 


DETERMINATION    OF    CARBON    DIOXIDE    AND    OXYGEN 


329 


of  the  mercury  manometer  or  by  a  graduated  Marey's  tambour.  Connect 
a  piece  of  gas  tubing  with  the  proximal  limb  of  the  mercury  manometer  and 
provide  it  with  a  glass  mouthpiece.  Insert  this  mouthpiece  well  back  into 
the  cavity  of  the  mouth,  closing  the  lips  firmly  about  it,  keeping  the 
pharynx  relaxed.  Note  the  variations  in  pressure  at  the  height  of  ordinary 
inspiration  and  expiration  through  the  nasal  passages.  Repeat  with 
forced  inspiration  and  expiration,  close  the  nasal  passages,  and  make  the 


FIG.  246. — Chart  showing  transverse  section  of  the  chest  at  the  level  of  the 
mamma  of  a  distance  runner,  age  twenty-one,  height,  5  feet,  n  inches,  net  weight,  140 
T^nnrie  ]?xi ?  ordinary  expiration;  Ex*,  forced  expiration;  In1,  ordinary  inspiration; 


pounds. 

In2,  forced  inspiration. 


Scale  J. 


maximal  expiratory  and  inspiratory  effort.     The  manometer  may  be  ad- 
justed to  write  on  the  smoked  paper  or  read  directly. 

7.  Demonstration    of    Carbon    Dioxide    in    Expired    Air. — Arrange 
two  flasks,  as  in  figure  246,  filling  each  one-third  full  of  baryta-water,  or 
lime-water.     Close  the  lips  around  the  mouthpiece  of  the  apparatus  and 
inhale  and  exhale  the  air  through  it.     The  inspired  air  will  come  through 
a,  the  expired  air  out  through  b.     The  baryta  water  in  b  will  quickly 
become  clouded  with  a  white  precipitate  of  barium  carbonate  while  that 
in  a  will  remain  clear  or  only  very  slightly  clouded,  showing  the  excess  of 
carbon  dioxide  in  expired  air. 

8.  Quantitative  Determination  of  Carbon  Dioxide  and  Oxygen  in 
Inspired  Air  and  in  Expired  Air. — Inspired  Air. — Fill  a  gas-analysis 
apparatus,  the  Guthrie  or  any  modern  modification  of  the  Haldane  or  the 
Orsat  analyzer,  with  air  from  outside  the  laboratory.     Read  the  volume 


330 


RESPIRATION 


at  room  temperature  and  pressure.  Wash  the  sample  back  and  forth 
through  the  potash  bulb  as  directed  ten  times  and  read  for  absorption  of 
the  carbon  dioxide.  Next  wash  in  ten  per  cent,  pyrogallic  acid  in  potas- 
sium hydroxide  until  constant  readings  show  that  all  oxygen  is  absorbed. 


FIG.  246a. — Apparatus  for  Demonstrating  Excess  of  CO2  in  Expired  Air.     Flasks  filled 

with  lime-water. 

The   nitrogen   residue   is   calculated  by   difference.     Compute   the  per- 
centages of  carbon  dioxide,  oxygen  and  nitrogen. 

Expired  Air. — Exhale  ten  expirations  controlled  by  one-way  valves 
into  an  ordinary  respirometer  or  a  Tissot  apparatus.  Now  fill  the  Guthrie 
apparatus  with  a  sample  of  this  expired  air  and  analyze  as  before,  first  for 


FIG.  2466. — Change  in  Respiration  on  Stimulating  the  Central  End  of  the  Sciatic 
Nerve.  The  rate  is  sharply  increased  and  the  amplitude  more  than  doubled.  The 
stimulation  is  between  the  points  marked  on  and  off,  time  in  seconds.  The  inspiratory 
movement  following  the  stimulation  was  greater  than  the  limit  of  the  recording  tambour. 

carbon  dioxide,  then  for  oxygen;  compute  the  percentage  of  each  gas, 
including  nitrogen.  The  expired  air  will  usually  be  found  to  have  lost 
from  4  to  5  per  cent,  of  oxygen  and  have  gained  a  little  more  than  that 
quantity  of  carbon  dioxide. 


THE    NERVOUS    MECHANISM    OF    RESPIRATORY    MOVEMENTS      331 


From  the  percentages  obtained  in  these  experiments,  and  the  volume 
of  air  breathed  per  unit  of  time,  computed  in  Experiment  4  above,  deter- 
mine the  amount  of  carbon  dioxide  exhaled  per  hour  per  kilogram  of 
weight  for  your  own  body.  Compute  also  the  amount  of  oxygen  con- 
sumed per  kilo  per  hour;  per  square  meter  of  surface  per  hour. 

9.  The  Rate  and  Character  of  the  Respiratory  Movements  in  the 
Mammal. — a,  The  rate  of  respiration  can  be  best  determined  by  direct 
count  per  minute,  an  effort  being  made  to  maintain  as  nearly  normal 


CK 


c  d 

FIG.  24.6c. — Oxygen  and  carbon   dioxide   analyzer,    Guthrie  form. 
FIG.  246^. — Receiver  for  air  sample,   Guthrie  form. 

conditions  as  possible.  Make  the  determinations  on  a  cat,  a  dog,  and 
a  guinea-pig,  b,  The  character  of  the  respiratory  movements  can  be 
recorded  by  one  of  the  various  forms  of  stethograph  adapted  to  the  size  of 
the  animal,  or  by  the  arrangement  shown  in  figure  233.  It  is  necessary  to 
make  the  determination  with  an  animal  under  the  influence  of  an  anes- 
thetic. 

10.  The  Nervous  Mechanism  of  Respiratory  Movements. — a.  The 
Ejffett  of  Stimulating  Cutaneous  Nerves. — Use  a  small  dog  or  a  cat  for 
this  experiment;  anesthetize  and  introduce  a  tracheal  tube  with  a  side 


332  RESPIRATION 

branch  adapted  for  measuring  the  variations  of  pressure  during  respira- 
tion. Connect  the  free  limb  of  the  tracheal  tube  with  an  ether  apparatus 
and  adjust  to  secure  constant  anesthesia.  Connect  the  side  branch  of  the 
tracheal  tube  with  a  Marey's  recording  tambour  of  medium  size  and  sup- 
plied with  a  comparatively  delicate  membrane.  The  amplitude  of  the 
movements  of  the  tambour  may  be  regulated  by  a  screw  compress  on  a 
connecting  tube.  Arrange  an  induction  coil  with  platinum  electrodes  in 
the  usual  manner  for  stimulating  by  means  of  the  interrupted  current. 
Record  the  results  of  the  experiment  along  with  the  variations  of  blood- 
pressure  on  a  continuous-paper  kymograph;  the  instrument  should  be 
supplied  with  a  time  signal,  a  stimulating  signal,  etc. 

Now  stimulate  the  skin  of  the  abdominal  region,  the  groin,  with  a  com- 
paratively strong  medium  induction  current,  figure  2466.  Dissect  out  the 
sciatic  nerve,  cut  it,  stimulate  the  central  end  with  a  mild  to  medium 
strength  of  current.  The  stimulus  should  be  graduated  carefully,  for 
there  is  often  such  a  great  increase  in  respiratory  rate  and  volume  that  the 
animal  may  become  overanesthetized. 

b.  The  Effect  of  Stimulating  the  Vagus  Nerve. — Isolate  and  stimulate 
the  vagus  nerve  with  a  medium  strength  of  stimulus.     The  effect  is  usually 
complete  inhibition  of  respiratory  movements.     By  means  of  graduated 
stimuli  one  may  demonstrate  the  accelerator  effects  from  the  stimulation 
of  the  vagus.     Stimulate  also  the  superior  laryngeal,  and  compare  with  the 
effects  of  stimulating  the  whole  vagus. 

c.  The  Effect  of  Cutting  the  Vagus  Nerves. — Isolate  both  vagus  nerves 
and  section  them  as  nearly  at  the  same  moment  as  possible.     Be  sure  to 
mark  on  the  tracing  the  exact  moment  at  which  the  nerves  are  cut.     This 
experiment  should  be  performed  with  every  accessory  condition  as  con- 
stant as  possible,  and  the  animal  should  not  be  disturbed  for  one  or  two 
minutes  so  that  the  effects  of  the  section  will  be  recorded,  figure  240.     The 
result  is  always  a  marked  deepening  and  slowing  of  the  respiratory 
movements. 

d.  The  Effect  of  Stimulating  the  Central  End  of  the  Vagus. — Upon  stimu- 
lating the  central  end  of  the  vagus  after  section,  the  respiration  rate  will  be 
inhibited  as  in  b,  showing  that  the  vagus  nerves  carry  afferent  respiratory 
fibers,  figure  241. 

e.  The  Effect  of  Stimulation  of  the  Phrenic  Nerves. — Isolate  the  right 
phrenic  nerve  at  its  origin  from  the  brachial  plexus  and  stimulate  it  with  a 
medium  strength  of  stimulus.     Upon  stimulating  the  nerve  the  diaphragm 
will  remain  in  contraction  and  the  record  will  show  that  the  thorax  is  in  the 
inspiratory  phase.     The  phrenic  is  therefore  a  simple  motor  nerve. 

Section  this  nerve  and  note  the  change  in  the  character  of  respiratory 
movements;  make  direct  observations  on  the  diaphragm,  examining  from 
the  abdominal  side. 


DEMONSTRATION    OF    APNEA,    DYSPNEA,    AND    ASPHYXIA        333 

ii.  Demonstration  of  Apnea,  Dyspnea,  and  Asphyxia. — Produce 
deep  anesthesia,  then  disconnect  the  ether  bottle  and  connect  the  tracheal 
tube  with  a  hand  bellows.  Produce  deep  and  forced  artificial  respiration 
for  ten  to  twenty  seconds.  Stop  artificial  respiration;  the  animal  will 
remain  quiet  and  without  any  effort  at  breathing.  This  is  the  condition 
of  apnea.  Allow  the  animal  to  recover  its  normal  respiratory  rate  and 
again  produce  deep  anesthesia.  Now  clamp  off  the  tracheal  tube  so  that 
the  animal  can  no  longer  receive  air  and  leave  it  so  until  death.  As  the 
blood  becomes  more  and  more  venous  there  will  first  be  a  marked  increase 
in  the  respiration  rate  and  depth.  This  is  known  as  hyperpnea.  This 
stage  is  followed  by  one  of  increasing  respiratory  amplitude  in  which  the 
accessory  respiratory  muscles  not  previously  active  are  brought  into 
forcible  contractions,  both  inspiratory  and  expiratory  phases  are  now 
forced,  dyspnea.  The  movements  continue  to  increase,  and  the  muscles 
of  the  neck,  larynx,  mouth,  and  nostrils  now  take  part.  There  is  a 
rather  sudden  decrease  in  the  respiratory  movements,  an  extension  of  the 
limbs,  and  gasping  movements,  after  which  the  animal  remains  quiet, 
death  being  produced  by  asphyxia. 

12.  Respiratory  Exchange  and  Calorimetry. — Indirect  calorimetry  is 
made  the  basis  for  determining  the  metabolic  rate  in  man  and  mammals. 
Measurement  of  the  oxygen  intake  and  of  the  carbondioxide  output  under 
standard  conditions  suffices  for  the  computation  from  the  data  of  the 
metabolic  rate  of  heat  production  per  kilo  or  per  square  meter  of  surface 
per  hour.  This  principle  is  used  in  the  rapid  methods  of  human  clinical 
calorimetry. 

Generally  a  man  is  made  to  exhale  into  a  large  collecting  chamber 
(Tissot  apparatus),  or  breathe  through  a  closed  chamber  (rebreather) 
provided  with  absorbers  and  either  one-way  valves  or  a  pump  to  circulate 
the  air  of  the  chamber. 

In  man  the  test  is  made  with  the  body  at  maximum  rest.  Test  (i) 
after  a  12  hour  fast;  (2)  measure  the  height  in  centimeters  and  weight 
in  kilos;  (3)  provide  20  to  30  minutes  complete  relaxation  lying  at  rest; 
(4)  measure  systolic  and  diastolic  blood  pressures  and  mouth  tempera- 
ture after  the  rest  period;  (5)  Adjust  the  face  mask  and  run  the  test  of 
oxygen  consumption.  Use  a  modified  Henderson  rebreather  or  a  Bene- 
dict filled  with  excess  of  pure  oxygen,  or  a  Tissot  apparatus.  Continue 
the  test  for  ten  minutes  or  more.  In  the  dog  use  a  rebreather  type  of 
apparatus  of  adapted  size.  Attach  a  face  mask  to  a  ten  kilo  dog  trained 
to  the  experiment.  Record  all  rebreather  movements. 

From  the  data  for  man,  compute  the  metabolic  rate  per  square  meter 
per  hour  for  man  by  DuBois'  formula. 

In  man  the  surface  area  in  sq.  cm.  =  The  Weight  in  kilos  °-425  X  the 
Height  in  cm.  °-725  X  a  constant  71.84. 


334  RESPIRATION 

Compute  the  rate  per  kilo  per  hour  for  the  dog.  Compare  with  the 
rate  per  kilo  on  man  and  explain  the  difference  observed. 

13.  Gases  of  Arterial  and  Venous  Blood. — Tissue  respiration  is  inti- 
mately dependent  on  the  capacity  of  the  blood  to  carry  oxygen  and  carbon 
dioxide  by  chemical  fixation,  chiefly  by  the  respiratory  pigment  hemo- 
globin. Measure  the  gaseous  content  of  arterial  and  venous  blood 
simultaneously  drawn  from  a  dog  under  local  analgesia.  Draw  the 
samples  under  petrolatum  and  analyze  promptly  for  the  three  gases, 
carbon  doxide,  oxygen  and  nitrogen,  in  a  Van  Slyke  apparatus,  page  303. 
Also  consult  the  Journal  of  Biological  Chemistry,  vol.  30,  page  347,  and 
vol.  33,  page  127. 


CHAPTER  VII 
SECRETION  IN  GENERAL 

ALL  tissues  of  the  body  produce  certain  chemical  changes  as  a  result  of 
their  protoplasmic  activity.  But  in  certain  cells  chemical  elaborations  have 
come  to  be  the  chief  function,  the  cells  have  been  differentiated  in  that  direc- 
tion, and  the  name  secreting  tissue  or  gland  tissue  is  applied.  The  end  result 
of  metabolism  in  gland  tissue  is  the  extrusion  on  the  free  borders  of  the  cells 
of  the  products  of  their  metabolism.  The  products  are  known  as  secretions 
and  the  process  itself  is  the  act  of  secretion.  Certain  secretions  which  are 
in  the  nature  of  waste  products  to  the  body  as  a  whole,  such  as  urine  in  the 
kidney,  are  often  called  excretions,  but  the  use  of  the  term  should  not  be  allowed 
to  confuse  the  general  similarity  of  this  to  other  secretions  as  regards  the 
physiological  changes  involved  in  its  production. 

Most  secretions  accomplish  some  definite  office  in  the  economy  of  the 
body.  Those  that  are  discharged  on  some  free  mucous  surface,  as  the  saliva, 
gastric  juice,  tears,  etc.,  are  called  external,  or  true  secretions,  or  merely  secre- 
tions. Substances  that  are  discharged  back  into  the  blood  stream  later  to 
influence  the  metabolism  of  tissues  other  than  the  ones  which  produced  them 
are  called  internal  secretions. 

Gland  cells,  like  other  tissues,  draw  their  nourishment  from  the  blood 
and  lymph.  The  product  or  secretion  of  gland  cells  may,  in  fact  usually 
does,  contain  some  of  the  substances  found  in  the  blood,  but  there  are  also 
present  new  materials  elaborated  by  the  cells,  and  even  where  the  same  sub- 
stance exists  both  in  the  secretion  and  in  the  blood  and  lymph  it  can  make 
its  appearance  in  the  secretion  only  under  the  control  of  the  protoplasm  of 
the  gland  cells.  The  saliva  secreted  by  the  salivary  cells,  for  example,  con- 
sists of  about  98  to  99  per  cent,  water  containing  in  solution  small  quantities 
of  certain  salts,  also  found  in  the  lymph,  and  a  small  percentage  of  the  en- 
zyme, ptyalin.  This  enzyme  is  peculiar  to  the  salivary  secretion  and  is  manu- 
factured by  the  salivary-cell  protoplasm.  As  is  well  known,  it  acts  vigorously 
in  extreme  dilution,  hence  the  high  per  cent,  of  water  in  the  secretion.  The 
passage  of  water  from  a  solution  as  concentrated  as  blood  plasma  to  a  solu- 
tion as  dilute  as  saliva  requires  a  high  amount  of  osmotic  energy,  an  amount 
that  can  be  supplied  only  from  the  chemical  energy  liberated  by  the  cell  in 
its  protoplasmic  activity.  After  the  removal  of  the  special  organ  by  which 
each  secretion  is  manufactured,  the  secretion  is  no  longer  formed.  Cases 
sometimes  occur  in  which  the  secretion  continues  to  be  formed  by  the  natural 

335 


336  SECRETION    IN    GENERAL 

organ,  but,  not  being  able  to  escape  toward  the  exterior,  on  account  of  some 
obstruction,  is  reabsorbed  and  accumulates  in  the  blood.  It  may  be  dis- 
charged from  the  body  in  other  ways;  but  these  are  not  instances  of  true 
vicarious  secretions,  and  must  not  be  so  regarded. 

Organs  and  Tissues  of  Secretion. — The  principal  secreting  organs 
are  the  following:  i.  The  serous  and  synovial  membranes;  2.  the  mucous 
membranes  with  their  special  glands,  e.g.,  the  buccal,  gastric,  and  intestinal 
glands;  3.  the  salivary  glands  and  pancreas;  4.  the  liver;  5.  the  mam- 
mary glands;  6.  the  lachrymal  glands;  7.  the  kidney  and  skin;  8.  the 
testes  and  ovaries,  and  9.  thyroid,  supra-renal,  etc. 

The  special  structure  and  functions  of  the  secreting  organs  will  be  given 
in  greater  detail  in  the  chapters  which  immediately  follow.  The  general 
types  of  structure  and  general  conditions  that  influence  the  functions  are 
introduced  at  this  point. 

Structural  Types  of  Secreting  Organs. — Serous  and  Synovial  Type.— 
The  serous  membranes  form  closed  sacs  lining  visceral  cavities  like  the 
abdominal,  pericardial,  or  pleural  cavities.  The  organs  are,  as  it  were, 
pushed  into  this  sac  and  carry  before  them  an  investment  of  membrane.  The 
serous  membranes  consist  of  a  single  layer  of  flattened  polygonal  cells 
resting  on  a  supporting  membrane  of  connective  tissue,  supporting  a  rami- 
fication of  blood  vessels,  lymphatics,  and  nerves. 

In  some  instances,  i.e.,  synovial  membranes,  the  secreting  layer  is  in- 
creased by  finger-like  elevations.  This  type  of  secreting  organ  produces 
ordinarily  only  enough  secretion  to  keep  the  surface  moist. 

The  Mucous  Type. — The  mucous  tracts,  and  different  portions  of  each 
of  them,  present  certain  structural  peculiarities  adapted  to  the  functions 
which  each  part  has  to  discharge;  yet  in  some  essential  characters  the  mucous 
membrane  is  the  same,  from  whatever  part  it  is  obtained.  In  all  the  princi- 
pal and  larger  parts  of  the  several  tracts  it  presents  an  external  layer  of  epithe- 
lium, situated  upon  a  basement  membrane,  and  beneath  this  a  stratum  of 
vascular  tissue  of  variable  thickness,  containing  lymphatic  vessels  and  nerves. 
The  vascular  stratum,  together  with  the  basement  membrane  and  epithelium, 
in  certain  cases  is  elevated  into  minute  papillae  and  villi,  in  others  depressed 
into  involutions  in  the  form  of  glands.  But  in  the  invaginations  of  the  secret- 
ing membrane  of  gland  cells,  the  supporting  basement  membrane  and  the  net- 
work of  capillaries  are  still  retained  in  their  relative  position.  With  increas- 
ing complexity  of  involution  the  simple  mucous  membrane  becomes  packed 
away  in  an  apparently  solid  mass.  The  equivalent  of  a  large  amount  of 
secreting  surface  is  thus  condensed  into  a  small  space.  In  the  process  of  in- 
vagination  some  differentiation  occurs  in  that  certain  of  the  gland  cells  be- 
come conducting  and  have  their  secretory  activity  somewhat  reduced. 

Secreting  Glands. — The  secreting  glands  present,  amid  manifold 
diversities  of  form  and  composition,  a  general  plan  of  structure;  but  all  are 


SECRETING    GLANDS 


337 


constructed  with  particular  regard  to  the  arrangement  of  the  cells  which  has 
just  been  described. 

Secreting  glands  are  classified  according  to  certain  structural  types,  as: 
i.  The  simple  tubular  gland,  A,  figure  247,  examples  of  which  are  furnished 
by  the  follicles  of  Lieberkuhn,  and  the  tubular  peptic  glands  of  the  stomach. 
They  are  simple  tubes  of  mucous  membrane,  the  walls  of  which  are  lined  with 


FIG.  247. — Plans  of  Extension  of  Secreting  Membrane  by  Inversion  or  Recession  in  the 
Forms  of  Cavities.  A,  Simple  glands,  viz.,  g,  straight  tube;  ht  sac;  i,  coiled  tube.  B, 
Multilocular  crypts;  k,  of  tubular  form;  /,  saccular.  C,  Racemose  or  saccular  compound 
gland;  m,  entire  gland,  showing  branched  duct  and  lobular  structure;  «,  a  lobule,  detached 
with  o,  branch  of  duct  proceeding  from  it.  D,  Compound  tubular  gland.  (Sharpey.) 

secreting  cells  arranged  as  an  epithelium.     To  the  same  class  may  be  re- 
ferred the  elongated  and  tortuous  sudoriferous  glands. 

2.  The  compound  tubular  glands,  D,  figure  247,  form  another  division. 
These  consist  of  main  gland  tubes,  which  divide  and  subdivide.  Each  gland 
may  be  made  up  of  the  subdivisions  of  one  or  more  main  tubes.  The  ulti- 
mate subdivisions  of  the  tubes  are  sometimes  highly  convoluted.  They  are 


338  SECRETION   IN    GENERAL 

formed  of  epithelium  of  various  forms,  supported  by  a  basement  membrane. 
The  larger  tubes  may  have  an  outside  coating  of  fibrous  areolar  or  muscular 
tissue.  The  salivary  glands,  pancreas,  Brunner's  glands,  kidney,  testes,  with 
the  lachrymal  and  mammary  glands,  are  examples  of  this  type,  but  present 
more  or  less  marked  variations  among  themselves. 

3.  The  racemose  glands,  in  which  a  number  of  vesicles  or  acini  are  arranged 
in  groups  of  lobules,  C,  figure  247.  The  Meibomian  follicles  are  examples 
of  this  kind  of  gland.  There  seem  to  be  glands  of  mixed  character,  com- 
bining some  of  the  characters  of  the  tubular  with  others  of  the  racemose  type; 
these  are  called  tubulo-racemose  or  tubulo-acinous  glands.  The  acini  are 
formed  by  a  kind  of  fusion  of  the  walls  of  several  vesicles,  which  thus  combine 
to  form  one  large  cavity  with  recesses  lined  or  filled  with  secreting  cells.  The 
smallest  branches  of  the  gland-ducts  sometimes  open  into  the  centers  of  these 
cavities;  sometimes  the  acini  are  clustered  round  the  extremities  or  by  the 
sides  of  the  ducts;  but,  whatever  secondary  arrangement  there  may  be,  all 
have  the  same  essential  character  of  rounded  groups  of  vesicles  containing 
gland  cells,  and  opening  by  a  common  central  cavity  into  minute  ducts, 
which  in  the  large  glands  converge  and  unite  to  form  larger  and  larger 
branches,  and  at  length  one  common  trunk  which  opens  on  a  free  surface. 

The  Process  of  Secretion. — The  process  of  secretion  is  dependent 
upon  the  activity  of  the  secreting  cells.  In  the  case  of  the  water  and  salts  the 
physical  processes  of  filtration  and  diffusion  may  play  a  part. 

The  chemical  processes  constitute  the  process  of  secretion  properly  so 
called,  as  distinguished  from  mere  transudation  spoken  of  above.  In  the 
process  of  secretion  various  materials  which  do  not  exist  as  such  in  the  blood 
are  manufactured  by  the  agency  of  the  gland  cells,  using  as  a  nutrient  fluid 
the  blood  or,  to  speak  more  accurately,  the  lymph  which  fills  the  interstices 
of  the  gland  textures. 

Evidences  in  favor  of  this  view  are:  i.  That  gland  cells  are  constituents 
of  all  glands,  however  diverse  their  outer  forms  and  other  characters,  and 
they  are  placed  in  all  glands  on  the  surfaces  or  in  the  cavity  whence  the  secre- 
tion is  poured.  2.  That  certain  materials  of  secretions  are  visible  with  the 
microscope  in  the  gland  cells  before  they  are  discharged.  Thus,  granules 
probably  representing  the  precursors  of  the  ferments  of  the  pancreas  may 
be  discerned  in  the  cells  of  that  gland.  Granules  of  uric  acid  are  found  in 
the  cells  of  the  kidneys  of  birds  and  fish,  and  fatty  particles,  like  those  of  milk, 
in  the  cells  of  the  mammary  gland. 

Certain  secreting  cells,  like  the  cells  of  the  sebaceous  glands,  appear  to 
develop,  grow,  and  attain  their  individual  perfection  by  appropriating  nutri- 
ment from  the  fluid  exuded  by  adjacent  blood  vessels  and  building  it  up  so 
that  it  shall  form  part  of  their  own  substance.  In  this  perfected  state  the  cells 
subsist  for  some  brief  time  and  then  appear  to  dissolve,  wholly  or  in  part,  and 
yield  their  contents  to  the  peculiar  material  of  the  secretion.  The  changes 


INFLUENCE    OF    THE    NERVOUS    SYSTEM    ON    SECRETION          339 

which  have  been  noticed  from  actual  experiment  in  the  cells  of  the  salivary 
glands,  pancreas,  and  peptic  glands  will  be  described  more  fully  in  the 
chapter  on  Digestion. 

Discharge  of  secretions  from  the  glands  may  either  take  place  as  soon  as 
formed,  or  the  secretion  may  be  long  retained  within  the  gland  or  its  ducts. 
The  former  is  the  case  with  the  sweat  glands.  But  the  secretions  of  those 
glands  whose  activity  of  function  is  periodical  are  usually  retained  in  the  cells 
in  an  undeveloped  form  during  the  period  of  the  gland's  inaction. 

When  discharged  into  the  ducts,  the  further  course  of  secretions  is  affected: 

(1)  partly  by  the  pressure  from  behind;  the  fresh  quantities  of  secretion  pro- 
pelling those  that  were  formed  before.     In  the  larger  ducts,  its  propulsion  is 

(2)  assisted  by  the  contraction  of  the  walls.     All  the  larger  ducts,  such  as 
the  ureter  and  common  bile  duct,  possess  in  their  coats  plain  muscular  fibers; 
they  contract  when  irritated,  and  sometimes  manifest  peristaltic  movements. 
Rhythmic  contractions  in  the  pancreatic  and  bile  ducts  have  been  observed, 
and  also  in  the  ureters  and  vasa  deferentia.     It  is  probable  that  the  contrac- 
tile power  extends  along  the  ducts  to  a  considerable  distance  within  the  sub- 
stance of  the  glands  whose  secretions  can  be  rapidly  expelled.     Saliva  and 
milk,  for  instance,  are  sometimes  ejected  with  much  force. 

Circumstances  Influencing  Secretion. — The  principal  conditions 
which  influence  secretion  are  variations  in  the  quantity  of  blood  and  varia- 
tions in  nerve  impulses  passing  to  the  gland  cells  over  secretory  nerve  fibers. 

An  increase  in  the  quantity  of  blood  traversing  a  gland,  as  in  nearly  all 
the  instances  before  quoted,  coincides  generally  with  an  augmentation  of  its 
secretion.  Thus  the  mucous  membrane  of  the  stomach  becomes  florid  when, 
on  the  introduction  of  food,  its  glands  begin  to  secrete.  The  mammary 
gland  becomes  much  more  vascular  during  lactation.  All  circumstances 
which  give  rise  to  an  increase  in  the  quantity  of  material  secreted  by  an  organ 
produce,  coincidently,  an  increased  supply  of  blood.  But  we  shall  see  that  a 
discharge  of  saliva  may  occur  under  extraordinary  circumstances  without  in- 
crease of  blood  supply,  and  so  it  may  be  inferred  that  this  condition  of  in- 
creased blood  supply  is  not  absolutely  essential  to  the  immediate  formation 
of  secretion,  but  that  it  favors  the  prolonged  activity  of  glands. 

Influence  of  the  Nervous  System  on  Secretion. — The  process  of 
secretion  is  largely  regulated  through  the  nervous  system.  The  exact  mode 
in  which  the  influence  is  exhibited  must  still  be  regarded  as  somewhat  obscure. 
In  part  it  exerts  its  influence  by  increasing  or  diminishing  the  quantity  of 
blood  supplied  to  the  secreting  gland,  in  virtue  of  the  power  which  it  exercises 
over  the  contractility  of  the  smaller  blood  vessels.  It  also  has  a  more  direct 
influence,  as  is  described  at  length  in  the  case  of  the  submaxillary  gland,  upon 
the  secreting  cells  themselves.  This  may  be  called  trophic  influence.  Its 
influence  over  secretion,  as  well  as  over  other  functions  of  the  body,  may  be 
excited  by  causes  acting  directly  upon  the  nervous  centers,  upon  the  nerves 


340  SECRETION   IN    GENERAL 

going  to  the  Secreting  organ,  or  upon  the  nerves  of  other  parts.  In  the  latter 
case  a  reflex  action  is  produced.  Thus  the  impression  produced  upon  the 
sensory  nerves  by  the  contact  of  food  in  the  mouth  leads  to  afferent 
nerve  impulses  to  the  secretory  center  in  the  central  nervous  system,  im- 
pulses which  are  reflected  by  the  nerves  supplying  the  salivary  glands, 
and  produce,  through  these,  a  more  abundant  secretion  of  the  saliva. 

Through  the  nerves,  various  conditions  of  the  brain  also  influence  the 
secretions.  Thus,  the  thought  of  food  may  be  sufficient  to  excite  an  abun- 
dant flow  of  saliva.  And,  probably,  it  is  the  mental  state  which  excites  the 
abundant  secretion  of  urine  in  hysterical  paroxysms,  as  well  as  the  perspira- 
tions, and  occasionally  diarrheas,  which  ensue  under  the  influence  of  terror, 
and  the  tears  excited  by  sorrow  or  excess  of  joy.  The  quality  of  a  secretion 
may  also  be  affected  by  mental  conditions,  as  in  the  cases  in  which,  through 
grief  or  passion,  the  secretion  of  milk  is  altered,  and  is  sometimes  so  changed 
as  to  produce  irritation  in  the  alimentary  canal  of  the  child. 


CHAPTER  VIII 
FOOD  AND  DIGESTION 

THE  term  digestion  includes  those  changes  taking  place  in  the  body  which 
bring  the  materials  of  the  food  into  such  condition  that  they  may  be  taken  up 
by  the  blood  and  lymphatic  vessels  and  thus  rendered  available  for  the  metab- 
olism of  the  tissues.  In  the  process  the  foods  are  rendered  more  soluble 
and  more  diffusible.  Certain  bodies  which  are  already  soluble  and  diffusible 
are  converted  into  forms  which  are  more  available  for  the  tissues;  as  an  ex- 
ample, cane-sugar,  although  both  soluble  and  diffusible,  cannot  be  readily 
used  by  the  body  until  it  is  converted  from  a  disaccharide  to  a  monosaccha- 
ride.  In  fact,  few  of  the  food  materials  are  fit  for  immediate  use  when  taken 
into  the  body  and  are  therefore  practically  useless  until  digested. 

FOOD  AND  FOOD  PRINCIPLES. 

We  have  been  accustomed  to  classify  foods  into  certain  main  groups, 
chiefly  according  to  their  chemical  character,  as  follows: 

Proteins. — Such  as  albumin,  myosin,  gluten,  casein,  etc.;  gluco-protein, 
nucleoprotein,  etc.;  gelatin,  elastin,  etc.  These  furnish  nitrogen  in  avail- 
able form. 

Carbohydrates. — Such  aG  starch,  dextrose,  cane-sugar,  etc. 

Fats. — Such  as  tristearin,  tripalmitin,  triolein. 

Minerals. — The  various  salines  found  in  animal  and  vegetable  food. 

Water. 

The  classes  of  foods  just  enumerated  usually  exist  in  mixtures  rather  than 
in  simple  forms,  as,  for  example,  a  beef  roast  contains  a  representative  of  each 
of  the  five  classes  enumerated,  though  it  is  composed  chiefly  of  water,  pro- 
teins, and  fats.  The  human  body  is  capable  of  using  materials  of  a  great 
variety  of  forms,  but  most  of  these  have  the  foods  mixed  in  such  a  way  as  to  give 
representatives  of  each  of  the  classes  above  in  certain  general  proportions. 

Nitrogenous  Foods. — The  Flesh  of  Animals,  e.g.,  beef,  veal,  mutton, 
pork,  bacon,  ham,  chicken,  eggs,  milk,  etc.,  are  typical  nitrogenous  foods. 

Of  these,  beef  and  eggs  are  richest  in  nitrogenous  matters,  containing 
about  20  per  cent.  Mutton  contains  about  18  per  cent.,  veal  16 . 5,  and  pork 
10.  Beef  is  firmer,  more  satisfying,  and  is  supposed  to  be  more  strengthen- 
ing than  mutton,  whereas  the  latter  is  more  digestible.  The  flesh  of  young 
animals,  such  as  lamb  and  veal,  is  less  digestible  and  less  nutritious.  Pork 
contains  a  large  amount  of  fat  and  is,  therefore,  comparatively  indigestible. 


342 


FOOD   AND    DIGESTION 


PERCENTAGE  COMPOSITION  AND  FUEL  VALUE  PER  POUND  OF  SOME  COMMON 
FOOD  STUFFS.      (ATWATER  AND   BRYANT.) 


Water. 
Per 
cent. 

Pro- 
tein. 
Per 

cent. 

Fat. 
Per 
cent. 

Carbo- 
hy- 
drate. 
Per 
cent. 

Ash. 
Per 

cent. 

Fuel 
Value. 
Calories 
per 
Pound 

Meat  (beef  round) 

71    6 

22     6 

2     8 

i    3 

CAQ 

Meat  (pork  loin)  

52  .0 

16.6 

30  .  i 

i  .0 

3*y 

i,  580 

Fish  (king  salmon)  

63.6 

17.8 

17.8 

i,  080 

Eggs.  .  .    .        

77.7 

13.4 

IO  .  S 

I  .  O 

720 

Milk  (cow's) 

87  .0 

7  .  7 

4  .  O 

5.  o 

O  .  7 

72  «; 

Milk  (human) 

80    7 

2     O 

31 

6  o 

O     2 

Cheese  (American)           

31.6 

28.8 

7C  .  Q 

o  .  •? 

7  .4. 

2.0  <;  ^ 

Butter 

II     O 

I  .  O 

8  1;  o 

7  .O 

•2  60  <c 

Bread  (white)  

33  •  2 

IO  .  O 

I  .  7 

^•6 

I  .  O 

.2  <  < 

Bread  (corn) 

7.8  .  0 

7  •  0 

4  .  7 

46  .  3 

2  .  2 

20  e; 

Rice 

12     7. 

8    0 

O     3 

7O    O 

O    4 

6  10 

Oatmeal  .        

7  .  1 

16.1 

7  •  2 

67.  < 

I  .  O 

860 

Beans  (dry) 

12.6 

22  .  <; 

1.8 

CQ  .  6 

7  .  e 

60  (? 

Potatoes  (white) 

78  1 

2     2 

O     I 

18  4 

I     O 

i8«; 

Potatoes  (sweet)  .           .           . 

60  .  o 

1.8 

o  .  7 

27    .4. 

I  .  2 

^70 

Fruit  (strawberries)  

GO  .4 

I  .  O 

0.6 

7.4 

0.6 

180 

Watermelon  (edible  portion)  

92.4 

0.4 

0  .  2 

6.7 

o-3 

140 

Meat  contains:  (i)  Muscle  proteins,  chiefly  myosin,  blood  proteins,  colla- 
gen (from  the  interstitial  fibrous  connective  tissue),  elastin  (from  the  elastic 
tissue),  as  well  as  traces  of  hemoglobin.  (2)  Fats,  including  the  lipoids 
lecithin  and  cholesterol.  (3)  Extractives,  some  of  which  are  agreeable  to 
the  palate  and  others  weakly  stimulating.  These  are  divided  into  the 
non-nitrogenous:  glycogen,  dextrose,  lactic  acid,  inosit,  etc.,  and  into  the 
nitrogenous:  consisting  chiefly  of  creatin,  and  the  purine  bases.  (4)  Salts, 
chiefly  chlorides  and  phosphates  of  potassium,  calcium,  and  magnesium. 
(5)  Water,  the  amount  of  which  varies  from  15  per  cent,  in  dried  bacon  to 
39  in  pork,  51  to  53  in  fat  beef  and  mutton,  and  72  per  cent,  in  lean  beef  and 
mutton. 
TABLE  OF  PERCENTAGE  COMPOSITION  OF  BEEF,  MUTTON,  PORK,  AND  VEAL. 

(L/ETHEBY.) 


Water. 

Protein. 

Fats. 

Salts. 

Beef  —  lean     ...      .           .  .           ... 

72 

IQ  .  1, 

3.6 

C.  I 

Beef  —  fat 

^1 

14.8 

20.8 

4  .4 

Mutton  —  lean  

72 

18.3 

4.9 

4.8 

Mutton  —  -fat  .... 

C7 

12  .4 

31.1 

7  .  C 

Veal 

6*. 

16.5 

15.8 

4  •  7 

Pork—  fat   . 

3Q 

0.8 

48.0 

2.3 

NITROGENOUS    FOODS  343 

TABLE  OF  PERCENTAGE  COMPOSITION  OF  POULTRY  AND  FISH.     (LETHEBY.) 


Water. 

Protein. 

Fats. 

Salts. 

Poultry  

74 

2  T  .  O 

1.8 

I  .  2 

White  fish  .  . 

78 

18.1 

2  .  0 

I  .  O 

Salmon 

16.  i 

5.  c 

I     4 

Eels  (very  rich  in  fat)  
Oysters.  . 

75 

7^.74. 

9.9 

II  .72 

13-8 
2  .42 

1.3 

2.73 

The  flesh  of  nearly  all  animals  has  been  occasionally  eaten,  and  we  may 
presume  that  except  for  difference  of  flavor,  etc.,  the  average  composition, 
aside  from  the  fat,  is  nearly  the  same  in  most  cases. 

Milk. — Milk  is  the  entire  food  of  young  animals,  and  contains  all  the 
elements  of  a  typical  diet.  Albuminous  substances  are  represented  in  the 
form  of  ca.seinogen,  and  serum  or  lactalbumin;  fats  in  thecream;  carbohydrates 
in  the  form  of  lactose  or  milk-sugar;  salts,  chiefly  as  calcium  phosphate;  and 
water.  From  milk  we  obtain  a  number  of  food  preparations,  such  as  cheese 
rich  in  protein  and  fat,  butter  and  cream,  buttermilk  rich  in  proteins  and 
peculiarly  well  adapted  for  invalid  diet,  and  whey  which  contains  all  the  sugar, 
salts  and  the  albumin. 

TABLE  OF  COMPOSITION  OF  MILK,  BUTTERMILK,  CREAM,  AND  CHEESE. 
(LETHEBY  AND  PAYEN.) 


Nitrogen- 
ous 
matters. 

Fats. 

Lactose. 

Salts. 

Water 

Milk  (cow}                           .  . 

4  •  * 

7  .  Q 

C  .  2 

0.8 

86 

Buttermilk 

4  .  I 

O  .  7 

6.4 

0.8 

88 

Cream 

4     I 

26    7 

2.8 

1.8 

66 

Cheese  —  ski  m 

44    8 

6   ^ 

4Q 

44 

Cheese  —  cheddar 

28.4 

3  1  •  i 

4  .  ^ 

^6 

Eggs. — The  yolk  and  albumin  of  eggs  of  oviparous  animals  bear  the 
same  relation  as  food  for  the  embryos  that  milk  bears  to  the  young  of  mam- 
malia, and  affords  another  example  of  the  natural  admixture  of  the  various 
alimentary  principles.  The  proteins  of  eggs  are  ovalbumin  and  ovoglobulin 
and  phosphoprotein,  the  mtellin  of  the  yolk.  In  addition  to  the  three  com- 
mon fats  there  is  a  yellow  fatty  pigment,  lutein  (lipochrome),  lecithin,  and 
cholesterol,  a  small  quantity  of  dextrose,  and  inorganic  salts,  chiefly  calcium, 
potassium,  sodium,  chlorides,  and  phosphates. 


344  FOOD    AND   DIGESTION 

TABLE  OF  THE  PERCENTAGE  COMPOSITION  OF  FOWLS'  EGGS. 


Nitrogenous 
substances. 

Fats. 

Salts. 

Water. 

White  

2O    4. 

i    6 

78 

Yolk  

16 

•2Q      7 

i    3 

r  2 

Legumes  are  used  by  vegetarians  as  the  principal  source  of  the  nitrogen  of 
the  food.  Those  chiefly  used  are  peas,  beans,  lentils,  etc.;  they  contain  a 
nitrogenous  substance  called  legumin,  allied  to  albumin.  Legumes  contain 
about  25.30  per  cent,  of  this  nitrogenous  body  and  twice  as  much  nitrogen 
as  wheat.  Nuts  also  form  a  very  nutritious  article  of  diet. 

Carbohydrate  Foods. — Bread,  made  from  ground  grain  obtained  from  the 
various  so-called  cereals,  viz.,  wheat,  rye,  maize,  barley,  rice,  oats,  etc.,  is 
the  direct  form  in  which  the  carbohydrate  is  supplied  in  an  ordinary  diet. 
It  contains  starch,  dextrin,  and  a  little  sugar.  It  also  contains  gluten,  com- 
posed of  vegetable  proteins,  and  a  small  amount  of  fat. 

TABLE  OF  PERCENTAGE  COMPOSITION  OF  BREAD  AND  FLOUR. 


Nitrogenous 
matters. 

Carbo- 
hydrates. 

Fats. 

Salts. 

Water. 

Bread  

8.1 

r  i 

i  .6 

2  .  "? 

•3  7 

Flour.  .        .  . 

10   8 

70    8  ? 

2     O 

I     7 

I  r 

Various  articles  besides  bread  are  made  from  flour,  e.  g.,  spaghetti,  maca- 
roni, etc.  Dextrin  and  a  small  amount  of  dextrose  are  present  in  bread, 
particularly  in  the  crust. 

Vegetables,  especially  potatoes.  They  contain  starch  and  sugar.  In 
cabbage,  turnips,  etc.,  the  salts  of  potassium  are  abundant. 

Fruits  contain  sugar  and  organic  acids,  tartaric,  malic,  citric,  and  others. 

Sugar,  chiefly  saccharose,  used  pure  or  in  various  sweetmeats. 

Oils  and  Fats. — The  substances  supplying  the  oils  and  fats  of  the  food 
are  chiefly  butter,  bacon  and  lard,  suet  (beef  and  mutton  fat),  and  vegetable 
oils.  These  contain  the  fats  olein,  stearin,  and  palmitin.  Butter  contains 
some  tributyrin,  while  vegetable  oils,  as  a  rule,  contain  no  stearin. 

Mineral  or  Inorganic  Foods. — The  salts  of  the  food.  Nearly  all  the 
substances  in  the  preceding  classes  contain  a  greater  or  less  amount  of  the 
salts  required  in  food.  Green  vegetables  and  fruit  contain  certain  salts, 
chiefly  potassium.  Sodium  chloride  is  an  essential  food;  it  is  contained  in 


EFFECT  OF  COOKING  ON  FOODS  345 

nearly  all  solid  foods,  but  so  much  is  required  that  it  has  also  to  be  taken  as  a 
condiment.  Potassium  salts  are  found  in  muscle,  nerve,  and  in  meats 
generally,  and  in  potatoes  and  other  vegetables.  Calcium  salts  are  contained 
in  eggs,  blood  of  meat,  wheat,  and  vegetables.  Iron  is  contained  in  hemo- 
globin, in  milk,  eggs,  and  vegetables. 

Liquid  Foods. — Water  is  essential  to  life,  and  from  two  to  two  and  a 
half  pints  a  day  must  be  consumed  in  addition  to  that  taken  mixed  with  solid 
food.  Of  the  non-alcoholic  substances  which  may  be  adored  to  it  for  flavoring 
purposes,  such  as  tea,  coffee,  cocoa,  etc.,  the  last  can  alone  be  considered  to 
have  a  certain  food  value,  as  it  contains  fats,  albuminous  material,  and  starch, 
the  other  constituents  of  such  substances  being  a  volatile  oil,  an  alkaloid 
caffeine,  and  tannic  acid.  The  food  value  of  alcoholic  beverages,  which  has 
long  been  a  subject  of  controversy,  as  now  generally  agreed  is  but  slight. 
Beer,  wines,  and  spirits  contain  ethyl  alcohol,  the  amount  varying  from  i .  5 
to  4 . 5  per  cent,  in  beer  to  40  to  80  per  cent,  in  spirits. 

The  Effect  of  Cooking  on  Foods. — In  general  terms  cooking  may 
be  said  to  render  food  more  easily  digestible,  both  directly  and  indirectly, 
through  increased  palatability.  Subjecting  food  to  high  degrees  of  heat  also 
serves  to  kill  parasites,  such  as  trichinae  and  the  various  tapeworms,  which 
may  be  present  and  alive  in  raw  meats.  In  the  case  of  meats  various  methods 
of  cooking  are  employed.  In  roasting,  the  meat  in  bulk  is  subjected  to  a 
high  temperature  in  an  oven  for  a  short  time,  250°  C.  for  15  minutes,  followed 
by  a  somewhat  lower  temperature,  175°  C.,  until  the  cooking  is  completed. 
This  causes  a  coagulation  of  the  outer  layers  of  albumin  so  that  the  juices 
of  the  meat  are  retained  until  the  center  of  the  mass  is  cooked  to  the  stage 
desired,  i.  e-  raised  to  a  temperature  of  63°  to  65°  C-  when  medium  done. 
In  boiling,  the  meat  is  first  immersed  in  boiling  water  for  a  time  and  then 
the  cooking  continues  at  a  lower  temperature.  In  a  broth,  the  extractives 
may  be  obtained  by  heating  the  meat  in  water  for  a  long  period.  Such  a 
broth  contains  the  flavoring  and  the  stimulating  extracts  of  the  meat,  but 
is  of  only  slight  nutritive  value.  A  temperature  below  the  coagulation  point, 
at  60°  C.,  will  extract  more  nutritive  protein  substance.  For  small  pieces 
of  meat,  broiling  practically  serves  the  same  purpose  as  does  roasting  for 
larger  pieces.  Frying,  as  usually  employed,  is  the  least  serviceable  method 
of  preparation,  since  the  fat  or  other  oily  material  used  so  permeates  the 
food  as  to  render  it  difficult  of  penetration  by  the  digestive  juices. 

Cooking  produces  upon  vegetables  the  necessary  effect  of  rendering  them 
softer,  so  that  they  can  be  more  readily  broken  up  in  the  mouth.  It  also 
causes  the  starch  grains  to  swell  and  burst,  and  so  aids  the  digestive  fluids 
in  penetrating  into  their  substance.  The  albuminous  matters  are  coagulated, 
and  the  gummy,  saccharine,  and  saline  matters  are  removed.  The  con- 
version of  flour  into  dough  is  effected  by  mixing  it  with  water,  and  adding  a 
little  salt  and  a  certain  amount  of  yeast.  Yeast  consists  of  the  cells  of  an 


FOOD   AND    DIGESTION 

organized  ferment  (Torula  cerevisice)\  this  plant  in  its  growth  changes  by 
ferment  action  the  sugar  produced  from  the  starch  of  the  flour,  and  a  quantity 
of  carbon  dioxide  and  some  alcohol  is  formed.  The  gas  together  with  the 
action  of  heat  during  baking  causes  the  dough  to  rise,  and  the  gluten  being 
coagulated,  the  bread  sets  as  a  permanently  vesiculated  mass. 

THE  PROCESS  OF  DIGESTION. 

The  Enzymes. — The  digestive  process  involves  both  mechanical  and 
chemical  changes.  The  former  are  secured  by  the  crushing  and  grinding 
in  the  mouth,  together  with  the  mixing  and  kneading  that  come  from  the 
peristalses  of  the  stomach  and  intestine.  The  chemical  changes  are  the 
most  important  factors  of  the  digestive  process.  The  various  secretions  that 
are  poured  into  the  mouth,  stomach,  and  intestines  all  contain  substances 
which  react  on  the  foods  to  render  the  latter  more  soluble.  The  special 
agency  in  each  secretion  is  the  presence  of  representatives  of  the  chemical 
groups  known  as  enzymes.  These  enzymes,  or  unorganized  ferments,  are 
the  essential  factors  in  the  secretions  which  produce  the  chemical  changes 
in  the  foods.  Their  predominant  action  is  one  of  hydrolytic  cleavage;  that 
is,  the  substance  acted  upon  takes  up  water  and  then  splits  into  two  different 
substances,  usually  of  the  same  class.  The  chemical  nature  of  the  en- 
zymes is  as  yet  undetermined  because  of  the  difficulty  of  getting  absolutely 
pure  specimens.  Their  mode  of  action  is  at  present  regarded  in  the  nature 
of  catalysis.  That  is  to  say,  the  enzymes  by  their  presence  facilitate  reactions 
that  would  otherwise  take  place  but  very  slowly.  Practically  all  are  formed 
in  the  glands  as  zymogens,  which  bear  the  same  relation  to  enzymes  as 
fibrinogen  does  to  fibrin;  they  are  transformed  to  enzymes  by  the  proper 
stimulus,  but  never  exist  as  such  in  the  glands. 

Each  enzyme  has  a  special  point  of  temperature  at  which  it  acts  best,  and 
any  change  in  the  temperature  retards  its  action;  the  action  is  suspended  at 
a  definite  point  of  low  temperature,  but  the  enzyme  is  not  destroyed  by  cold. 
The  action  is  suspended  at  a  somewhat  higher  temperature,  and  at  a  still 
higher  point  the  enzyme  is  destroyed.  Some  enzymes  act  only  in  an  alka- 
line medium,  being  destroyed  in  an  acid  medium,  and  vice  versa.  Others 
act  in  either  alkaline,  or  neutral,  or  acid  media.  Enzymes  are  hindered  in 
their  action  by  the  accumulation  of  the  products  of  their  activity.  Most  of 
them  cease  acting  altogether  when  these  products  reach  a  certain  concen- 
tration, but  will  begin  acting  again  on  the  removal  of  these  products  or  if 
the  mixture  be  simply  diluted. 

The  quantity  of  the  enzyme  determines  the  rapidity  of  the  action,  but  not 
the  amount;  a  small  quantity  will  digest  as  much  as  a  large  quantity,  but  will 
take  longer.  The  enzymes  are  not  used  up  in  the  course  of  their  activity, 
as  far  as  can  be  seen,  and  do  not  seem  to  undergo  any  change  in  their 
composition. 


DIGESTION   IN    THE    MOUTH  347 

Enzymes  are  more  or  less  specific  in  their  action.  That  is,  each  enzyme 
is  supposed  to  produce  its  change  in  only  one  particular  substance,  as  in 
starch,  maltose,  protein,  fat,  etc.  An  enzyme  that  can  cause  cleavage  of  the 
starch  molecule  will  not  act  on  fat  or  protein  or  even  on  other  members  of 
the  starch  group.  This  specific  action  is  doubtless  expressive  of  a  definite 
relation  between  the  structure  of  the  enzyme  and  the  substance  acted  on. 

An  interesting  fact  as  to  enzyme  action  is  its  reversibility — a  phenomenon 
now  well  known  and  well  established  for  carbohydrates  and  fats.  Kastle 
and  Lowenhart  have  shown  that  lipase,  which  acts  to  split  neutral  fats  into 
fatty  acid  and  glycerin,  will  also  produce  a  synthesis,  at  least  of  butyric 
acid  and  alcohol  into  ethylbutyrate.  Taylor  and  Robertson  in  independent 
papers  have  recently  made  the  far-reaching  discovery  that  the  protein 
molecule  can  be  synthesized  by  the  agency  (apparent  reversible  action)  of 
enzymes. 

Enzymes  are  classified  either  according  to  the  chemical  nature  of  their 
action  or  according  to  the  class  of  substances  on  which  they  act;  the  former 
classification  is  more  logical,  but  the  latter  is  more  convenient  and  more 
generally  used. 

TABLE  OF  DIGESTIVE  ENZYMES. 

Amylolytic. 

Ptyalin  of  saliva,  and  amylopsin  of  pancreatic  juice,  change  starch  to  mal- 
tose.     Maltase  in   the   saliva,    and  pancreatic  juice  in   the   small   intestine, 
change  maltose  to  dextrose.     Lactase  splits  lactose  to  galactose  and  dextrose, 
and  invertase  splits  cane-sugar  to  levulose  and  dextrose  in  the  small  intestine. 
Li  poly  tic. 

Steapsin  or  lipase,  found  in  the  pancreatic  juice,  splits  neutral  fats   into 
glycerin  and  fatty  acid. 
Proteolytic. 

Pepsin  of  the  gastric  secretion,  and  trypsin  of  the  pancreatic  secretion, 
change  proteins  to  proteoses  and  peptones,  trypsin  breaking  the  protein  down 
to  simpler  nitrogenous  products.     Erepsin  of  the  intestine  splits  peptones  to 
simpler  products. 
Coagulating. 

Rennin  of  the  gastric  juice  coagulates  milk. 
Activating. 

Enterokinase  of  the  intestinal  juice  converts  trypsinogen  to  trypsin. 
(Thrombokinase  of  the  blood  is  of  this  class.) 

DIGESTION  IN  THE  MOUTH. 

The  food  is  received  into  the  mouth  and  is  subjected  to  the  action  of  the 
teeth  and  tongue,  being  at  the  same  time  mixed  with  the  first  of  the  digestive 
juices,  the  saliva.  It  is  then  swallowed,  and,  passing  through  the  pharynx 
and  esophagus  into  the  stomach,  is  subjected  to  the  action  of  the  gastric 
juice,  the  second  digestive  juice.  Thence  it  passes  into  the  small  intestines, 
where  it  meets  with  the  bile,  the  pancreatic  juice,  and  the  intestinal  juices,  all 


34-8  FOOD    AND    DIGESTION 

of  which  exercise  a  digestive  influence  upon  the  portion  of  the  food  not  already 
digested  and  absorbed.  In  the  large  intestine  some  further  digestion  and 
absorption  take  place,  and  the  residue  of  undigested  matter  leaves  the  body 
in  the  form  of  feces. 

Mastication. — The  act  of  mastication  is  performed  by  the  biting  and 
grinding  movement  of  the  lower  range  of  teeth  against  the  upper.  The 
simultaneous  movements  of  the  tongue  and  cheeks  assist  by  crushing  the 
softer  portions  of  the  food  against  the  hard  palate  and  gums,  thus  supple- 
menting the  action  of  the  teeth,  and  by  returning  the  morsels  of  food  to  the 
action  of  the  teeth  as  they  are  squeezed  out  from  between  them  until  they 
have  been  sufficiently  chewed. 

The  simple  up-and-down  or  biting  movements  of  the  lower  jaw  are  per- 
formed by  the  temporal,  masseter,  and  internal  pterygoid  muscles,  the  action 
of  which  in  closing  the  jaws  alternates  with  that  of  the  digastric  and  other 
muscles  passing  from  the  os  hyoides  to  the  lower  jaw,  which  open  the  jaws. 
The  grinding  or  side  movements  of  the  lower  jaw  are  performed  mainly  by 
the  external  pterygoid  muscles,  the  muscle  of  one  side  acting  alternately  with 
the  other.  When  both  external  pterygoids  act  together,  the  lower  jaw  is 
pulled  directly  forward,  so  that  the  lower  incisor  teeth  are  brought  in  front 
of  the  level  of  the  upper. 

The  act  of  mastication  is  voluntary.  It  will  suffice  here  to  state  that  the 
afferent  nerves  chiefly  concerned  are  the  sensory  branches  of  the  fifth,  ninth, 
and  tenth,  and  the  efferent  are  the  motor  branches  of  the  fifth  and  the  twelfth 
cerebral  nerves. 

The  act  of  mastication  is  much  assisted  by  the  saliva,  which  is  secreted  by 
the  salivary  glands  in  largely  increased  amount  during  the  process.  The 
intimate  incorporation  of  the  saliva  with  the  food  is  termed  insalivation. 

The  Salivary  Glands. — The  glands  which  secrete  the  saliva  in  the 
human  subject  are  the  salivary  glands  proper,  the  parotid,  the  submaxil- 
lary,  and  the  sublingual,  and  numerous  smaller  bodies  of  similar  structure 
and  with  separate  ducts,  which  are  scattered  thickly  beneath  the  mucous 
membrane  of  the  lips,  cheeks,  soft  palate,  and  root  of  the  tongue. 

Histological  Structure. — The  salivary  glands  are  compound  tubular 
or  tubulo-racemose  glands.  They  are  made  up  of  lobules.  Each  lobule  con- 
sists of  the  branchings  of  a  division  of  the  main  duct  of  the  gland,  which 
are  generally  more  or  less  convoluted  toward  the  extremities,  that  form  the 
alveoli,  or  proper  secreting  parts  of  the  gland.  The  salivary  secreting  cells 
are  of  cubical  or  columnar  form  and  are  arranged  around  a  central  canal. 
The  granular  appearance  frequently  seen  in  the  salivary  cells  is  due  to  the 
numerous  zymogen  granules  which  they  contain. 

During  the  rest  period  the  cells  are  larger,  highly  granular,  with  obscured 
nuclei  and  smaller  lumen.  During  activity  the  cells  become  smaller  and 
their  contents  more  opaque. 


NERVOUS    MECHANISM    OF    SECRETION    OF    SALIVA 


349 


When  the  mucous  type  of  gland  is  secreting,  or  on  stimulation  of  the  nerve, 
mucinogen  is  converted  into  mucin,  the  cells  swell  up,  appear  more  transparent 
and  stain  deeply  in  logwood,  figure  249.  After  stimulation,  the  cells  become 
smaller,  more  granular,  and  more  easily  stained  from  having  discharged  their 
contents,  and  the  nuclei  appear  more  distinct. 

Nerves  of  large  size  are  found  in  the  salivary  glands.  They  are  princi- 
pally contained  in  the  connective  tissue  of  the  alveoli,  and  certain  glands, 
especially  in  the  dog,  are  provided  with  ganglia.  Some  nerves  have  special 


FIG.  248. 


FIG.  249. 


FIG.  248. — Section  of  the  Submaxillary  Gland  of  a  Dog,  Resting  Stage.  Most  of  the 
alveolar  cells  are  large  and  clear,  being  filled  with  the  material  for  secretion  (in  this  case, 
mucigen),  which  obscures  their  protoplasm;  some  of  the  cells,  however,  are  small  and 
protoplasmic,  forming  the  crescents  seen  in  most  of  the  alveoli.  (Ranvier.) 

FIG.  249. — Section  of  a  Similar  Gland  after  a  Period  of  Activity.  The  mucigen  has 
been  discharged  from  the  mucin-secreting  cells,  which  consequently  appear  shrunken  and 
less  clear.  Both  the  cells  and  the  alveoli  are  much  smaller,  and  the  protoplasm  of  the  cells 
is  more  apparent.  The  crescents  of  Gianuzzi  are  enlarged,  c,  Crescent  cells;  g,  mucus- 
secreting  cells;  /,  lumen  of  alveolus.  (Ranvier.) 

endings  in  Pacinian  corpuscles,  some  supply  the  blood  vessels,  and  others 
penetrate  the  basement  membrane  of  the  alveoli  and  end  upon,  but  not  in, 
the  salivary  cells. 

The  blood  vessels  form  a  dense  capillary  network  around  the  ducts  of  the 
alveoli,  being  carried  in  by  the  fibrous  trabeculae  between  the  alveoli,  in  which 
also  the  lymphatics  begin  by  lacunar  spaces. 

The  Nervous  Mechanism  of  the  Secretion  of  Saliva. — The  secretion 
of  saliva  is  under  the  control  of  the  nervous  system.  Under  ordinary  con- 
ditions it  is  excited  by  the  stimulation  of  the  peripheral  branches  of  two 
nerves,  the  gustatory  or  lingual  branch  of  the  inferior  maxillary  division 
of  the  fifth  nerve,  and  of  the  glosso-pharyngeal,  which  are  distributed  to  the 
mucous  membrane  of  the  tongue  and  pharynx  conjointly.  The  stimulation 
occurs  on  the  introduction  of  sapid  substances  into  the  mouth,  and  the 
secretion  is  brought  about  in  the  following  way:  From  the  terminations  of 
the  above-mentioned  sensory  nerves  distributed  in  the  mucous  membrane 


350  FOOD   AND    DIGESTION 

an  impression  is  conveyed  upward  (afferent)  to  the  special  nerve  center 
situated  in  the  medulla  oblongata  which  controls  the  process,  and  by  it  is 
reflected  to  certain  nerves  supplied  to  the  salivary  glands,  which  will  be  pres- 
ently indicated.  In  other  words,  the  center,  when  stimulated  to  action  by 
the  sensory  impressions  carried  to  it,  sends  out  impulses  along  efferent  or 
secretory  nerves  supplied  to  the  salivary  glands.  These  cause  the  saliva  to  be 
secreted  by,  and  discharged  from,  the  gland  cells.  Other  stimuli,  however, 
besides  that  of  the  food,  and  other  sensory  nerves  than  those  mentioned 
may  reflexly  produce  the  same  effects.  For  example,  saliva  may  be  caused 
to  flow  by  irritation  of  the  mucous  membrane  of  the  mouth  with  mechanical, 
chemical,  electrical,  or  thermal  stimuli,  also  by  the  irritation  of  the  mucous 
membrane  of  the  stomach  in  some  way,  as  in  nausea  which  precedes  vomit- 
ing when  some  of  the  peripheral  fibers  of  the  vagi  are  irritated.  Stimulation 
of  the  olfactory  nerves  by  smell  of  food,  of  the  optic  nerves  by  the  sight  of  it, 
and  of  the  auditory  nerves  by  the  sounds  which  are  known  by  experience  to 
accompany  the  preparation  of  a  meal  may  also  stimulate  the  nerve  center  to 
action.  In  addition  to  these,  as  a  secretion  of  saliva  follows  the  movement 
of  the  muscles  of  mastication,  it  may  be  assumed  that  this  movement  stimu- 
lates the  secreting  nerve  fibers  of  the  gland,  direct  or  reflexly.  From  the  fact 
that  the  flow  of  saliva  may  be  increased  or  diminished  by  mental  states,  it 
is  evident  that  impressions  from  the  cerebrum  also  are  capable  of  stimulating 
the  center  to  action  or  of  inhibiting  its  action. 

Influence  of  Nerves  on  the  Submaxillary  Gland. — The  submaxillary 
gland  has  been  the  gland  chiefly  employed  for  the  purpose  of  experimentally 
demonstrating  the  influence  of  the  nervous  system  upon  the  secretion  of 
saliva,  because  of  the  comparative  facility  with  which  the  gland,  with  its  blood 
vessels  and  nerves,  can  be  exposed  to  view  in  the  dog,  rabbit,  and  other 
animals. 

The  chief  nerves  supplied  to  the  gland  are:  (i)  the  chorda  tympani,  a 
branch  given  off  from  the  facial  in  the  canal  through  which  it  passes  in  the 
temporal  bone;  and  (2)  branches  of  the  sympathetic  nerve  from  the  plexus 
around  the  facial  artery  and  its  branches  to  the  gland.  The  chorda,  figure 
250,  passes  downward  and  forward,  under  cover  of  the  external  ptery- 
goid  muscle,  and  joins  the  lingual  or  gustatory  nerve,  proceeds  with  it  for  a 
short  distance,  and  then  passes  along  the  sub  maxillary- gland  duct, 
giving  branches  to  the  submaxillary  ganglion,  and  sending  others  to 
terminate  in  the  superficial  muscles  of  the  tongue.  It  consists  of  fine  medul- 
lated  fibers  which  lose  their  medullae  in  the  gland.  If  this  nerve  be  exposed 
and  divided  anywhere  in  its  course  from  its  exit  from  the  skull  to  the  gland  no 
immediate  result  will  follow,  nor  will  stimulation  either  of  the  lingual  or  of 
the  glosso-pharyngeal  produce  a  flow  of  saliva.  But  if  the  peripheral  end 
of  the  divided  nerve  be  stimulated,  an  abundant  secretion  of  saliva  ensues, 
and  the  blood  supply  is  enormously  increased  by  dilatation  of  the  arteries. 


INFLUENCE    OF    NERVES    ON    THE    SUBMAXILLARY    GLAND         351 

The  veins  may  even  pulsate,  and  the  blood  contained  within  them  is  more 
arterial  than  venous  in  character. 

When,  on  the  other  hand,  the  stimulus  is  applied  to  the  sympathetic  fila- 
ments (mere  division  producing  no  apparent  effect),  the  arteries  contract, 
and  the  blood  stream  is  in  consequence  much  diminished;  and  only  a  sluggish 
stream  of  dark  blood  escapes  from  the  veins.  The  saliva,  instead  of  being 
abundant  and  watery,  becomes  scanty  and  tenacious.  If  both  chorda  tym- 
pani  and  sympathetic  branches  be  divided,  the  gland,  released  from  nervous 
control,  may  secrete  continuously  and  abundantly  (paralytic  secretion). 


FIG.  250. — Diagram  showing  the  distribution  of  the  cranial  and  sympathetic  secretory 
and  vase-motor  nerves  for  the  parotid  and  submaxillary  glands.  The  post-ganglionic 
neurones  are  in  black;  the  pre-ganglionic  neurones  including  the  central  neurone  of  the 
sympathetic  path  are  in  red.  (Diagram  based  on  figures  by  Sheldon,  Brubaker,  and 
Starling.) 


The  abundant  secretion  of  saliva  which  follows  stimulation  of  the  chorda 
tympani  is  not  merely  the  result  of  a  nitration  of  fluid  from  the  blood  vessels, 
in  consequence  of  the  largely  increased  circulation  through  them.  This  is 
proved  by  the  fact  that,  when  the  main  duct  is  obstructed,  the  pressure  within 
may  considerably  exceed  the  blood-pressure  in  the  arteries ;  and  also  that,  when 
some  atropine  has  been  previously  injected  into  the  veins  of  the  animal  ex- 
perimented upon  stimulation  of  the  peripheral  end  of  the  divided  chorda  pro- 
duces all  the  vascular  effects  as  before,  without  any  secretion  of  saliva  accom- 
panying them.  Again,  if  an  animal's  head  be  cut  off,  and  the  chorda  be 
rapidly  exposed  and  stimulated  with  an  interrupted  current,  a  secretion  of 


352  FOOD    AND    DIGESTION 

saliva  ensues  for  a  short  time,  although  the  blood  supply  is  necessarily  absent. 
These  experiments  serve  to  prove  that  the  chorda  contains  two  sets  of  nerve 
fibers:  one  set,  V  as  o- dilator ,  which,  when  stimulated,  act  upon  a  local  vaso- 
motor  center  for  regulating  the  blood  supply,  inhibiting  its  action,  and 
causing  the  vessels  to  dilate,  and  so  producing  an  increased  supply  of  blood 
to  the  gland;  while  another  set,  which  are  paralyzed  by  injection  of  atropine, 
directly  stimulate  the  cells  themselves  to  activity,  whereby  the  cells  secrete 
and  discharge  the  constituents  of  the  saliva  which  they  produce,  the  secretory 
nerves.  These  latter  fibers  very  possibly  terminate  on  the  salivary  cells 
themselves.  If,  on  the  other  hand,  the  sympathetic  fibers  be  divided,  stimu- 
lation of  the  tongue  by  sapid  substances,  or  electrical  stimulation  of  the  trunk 
of  the  lingual  or  of  the  glosso-pharyngeal,  continues  to  produce  a  flow  of 
saliva.  From  these  experiments  it  is  evident  that  the  chorda^  tympani  nerve 
is  the  principal  nerve  through  which  efferent  impulses  proceed  from  the  center 
to  excite  the  secretion  of  this  gland. 

The  sympathetic  nerve  also  contains  two  sets  of  fibers,  v  as  o- constrictor 
and  secretory.  But  the  flow  of  saliva  upon  stimulating  the  sympathetic  is 
scanty,  and  the  saliva  itself  viscid.  At  the  same  time  the  vessels  of  the  gland 
are  constricted.  The  secretory  fibers  may  be  paralyzed  by  the  administra- 
tion of  atropine. 

Nerves  of  the  Parotid  Gland. — The  nerves  which  influence  secretion 
in  the  parotid  gland  are  branches  of  the  facial  (lesser  superficial  petrosal) 
and  of  the  sympathetic.  The  former  nerve,  after  passing  through  the  otic 
ganglion,  joins  the  auriculo- temporal  branch  of  the  fifth  cerebral  nerve,  and, 
with  it,  is  distributed  to  the  gland.  The  nerves  by  which  the  stimulus  ordi- 
narily exciting  secretion  is  conveyed  to  the  medulla  oblongata  are,  as  in  the 
case  of  the  submaxillary  gland,  the  fifth  and  the  glosso-pharyngeal.  The 
pneumogastric  nerves  convey  a  further  stimulus  to  the  secretion  of  saliva 
when  food  has  entered  the  stomach;  the  nerve  center  is  the  same  as  in  the  case 
of  the  submaxillary  gland. 

Changes  in  the  Gland  Cells. — The  method  by  which  the  salivary 
cells  produce  the  secretion  of  saliva  appears  to  be  divided  into  two  stages, 
which  differ  somewhat  according  to  the  class  to  which  the  gland  belongs,  viz., 
whether  to  (i)  the  true  salivary  or  to  (2)  the  mucous  type.  In  the  former 
case  it  has  been  noticed,  as  already  described,  that  during  the  rest  which 
follows  an  active  secretion  the  lumen  of  the  alveolus  becomes  smaller,  the 
gland  cells  larger  and  very  granular.  During  secretion  the  alveoli  and  their 
cells  become  smaller,  and  the  granular  appearance  in  the  latter  to  a  consider- 
able extent  disappears,  and  at  the  end  of  secretion  the  granules  are  confined 
to  the  inner  part  of  the  cell  nearest  to  the  lumen,  which  is  now  quite  distinct, 
figure  251. 

It  is  supposed  from  these  appearances  that  the  first  stage  in  the  act  of 
secretion  consists  in  the  protoplasm  of  the  salivary  cell  taking  up  from  the 


CHANGES  IN  THE  GLAND  CELLS  353 

lymph  certain  materials  from  which  it  manufactures  the  elements  of  its  own 
secretion,  and  which  are  stored  up  in  the  form  of  granules  in  the  cell  during 
rest;  the  second  stage  consists  of  the  actual  discharge  of  these  granules,  with 
or  without  previous  change.  The  granules  are  zymogen  granules,  and  repre- 
sent the  chief  substance  of  the  salivary  secretion,  ptyalin.  In  the  case  of  the 
submaxillary  gland  of  the  dog,  at  any  rate,  the  sympathetic  nerve  fibers  ap- 
pear to  have  to  do  with  the  first  stage  of  the  process,  and  when  stimulated 
the  protoplasm  is  extremely  active  in  manufacturing  the  granules,  whereas 
the  chorda  tympani  is  concerned  in  the  production  of  the  second  act,  the 
actual  discharge  from  the  cells  of  the  materials  of  secretion,  together  with  a 


FIG.  251. — Alveoli  of  True  Salivary  Gland.     A,  At  rest;  B,  in  the  first  stage  of  secretion; 
C,  after  prolonged  secretion.     (Langley.) 

»• 

considerable  amount  of  fluid.  The  latter  is  an  actual  secretion  by  the 
protoplasm,  as  it  ceases  to  occur  when  atropine  has  been  subcutaneously 
injected. 

In  the  mucus-secreting  gland,  the  changes  in  the  cells  during  secretion 
have  been  already  spoken  of.  They  consist  in  the  gradual  production  by  the 
protoplasm  of  the  cell  of  a  substance  called  mucigen,  which  is  converted  into 
mucin,  and  discharged  on  secretion  into  the  canal  of  the  alveoli.  The  muci- 
gen  is,  for  the  most  part,  collected  into  the  inner  part  of  the  cells  during  rest, 
pressing  the  nucleus  and  the  small  portion  of  the  protoplasm  which  remains 
against  the  limiting  membrane  of  the  alveoli. 

The  process  of  secretion  in  the  salivary  glands  is  identical  with  that  of 
glands  in  general.  The  cells  which  line  the  ultimate  branches  of  the  ducts 
are  the  agents  by  which  the  special  constituents  of  the  saliva  are  formed.  The 
material  which  they  have  incorporated  within  themselves,  which  is  doubtless 
a  product  of  the  metabolism  of  the  protoplasm  of  the  cells,  is  given  up  again 
almost  at  once  in  the  form  of  a  fluid,  secretion,  which  escapes  from  the  ducts 
of  the  gland.  The  cells  themselves  undergo  diminution  in  the  mass  of  their 
protoplasm,  which  is  again  renewed  in  the  intervals  of  the  active  exercise  of 
the  functions.  The  source  whence  the  cells  obtain  the  materials  for  the  con- 
struction of  secretion  is  the  blood  plasma,  which  is  filtered  off  from  the  circu- 
lating blood  into  the  interstices  of  the  glands,  as  in  all  living  tissues. 


354  FOOD   AND    DIGESTION 

Saliva. — Saliva,  as  it  commonly  flows  from  the  mouth,  is  the  mixed 
secretion  of  the  salivary  glands  proper  and  of  the  glands  of  the  buccal  mucous 
membrane  and  tongue.  When  obtained  from  parotid  ducts,  and  free  from 
mucus,  saliva  is  a  transparent  watery  fluid,  the  specific  gravity  of  which 
varies  from  i .  004  to  i .  008  and  in  which,  when  examined  with  the  micro- 
scope, are  found  floating  a  number  of  minute  particles,  derived  from  the 
secreting  ducts  and  vesicles  of  the  glands.  In  the  impure  or  mixed  saliva 
are  found,  besides  these  particles,  numerous  epithelial  scales  separated  from 
the  surface  of  the  mucous  membrane  of  the  mouth  and  tongue,  and  the  so- 
called  salivary  corpuscles,  discharged  probably  from  the  mucous  glands  of  the 
mouth  and  the  tonsils.  These  subside  when  the  saliva  is  collected  in  a  deep 
vessel  and  left  at  rest.  They  form  a  white  opaque  sediment  leaving  the 
supernatant  fluid  transparent  and  colorless,  or  with  a  pale  bluish-gray  tint. 
Saliva  also  contains  various  kinds  of  micro-organisms  (bacteria).  The 
saliva,  when  first  secreted,  appears  to  be  always  alkaline  in  reaction;  the 
alkalinity  is  about  equal  to  o .  08  per  cent,  of  sodium  carbonate,  and  is  due 
to  the  presence  of  disodium  phosphate,  Na2HPO4. 

The  presence  of  potassium  sulpho cyanide,  KCNS,  in  saliva  may  be  shown 
by  the  blood-red  coloration  which  the  fluid  gives  with  a  solution  of  ferric 
chloride,  Fe2Cl6,  and  which  is  bleached  on  the  addition  of  a  solution  of 
mercuric  chloride,  HgCl2,  but  not  by  hydrochloric  acid. 

CHEMICAL  COMPOSITION  OF  HUMAN  SALIVA.      (HAMMERBACHER.) 

In  1,000  Parts. 

Water 994 .  2 

Solids 5.8 

Mucus  and  epithelium 2.2 

Soluble  organic  matter  (ptyalin) 1.4 

Potassium  sulphocyanide o .  04 

Salts .    2.20 

Saliva  from  the  parotid  is  less  viscid;  less  alkaline,  the  first  few  drops 
discharged  in  secretion  being  even  acid  in  reaction;  clearer,  although  it  may 
become  cloudy  on  standing  from  the  precipitation  of  calcium  carbonate  by 
the  escape  of  carbon  dioxide;  and  more  watery  than  that  from  the  submaxil- 
lary.  It  has,  moreover,  a  less  powerful  action  on  starch.  Sublingual  saliva 
is  the  most  viscid,  and  contains  more  solids  than  either  of  the  other  two,  but 
has  little  diastasic  action. 

Rate  of  Secretion  and  Quantity  of  Saliva. — The  rate  at  which  saliva 
is  secreted  is  subject  to  considerable  variation.  When  the  tongue  and  muscles 
concerned  in  mastication  are  at  rest,  and  the  nerves  of  the  mouth  are  subject 
to  no  unusual  stimulus,  the  quantity  secreted  is  not  more  than  sufficient  with 
the  mucus  to  keep  the  mouth  moist.  During  actual  secretion  the  flow  is 
much  accelerated. 


FUNCTION    OF    SALIVA 


355 


The  quantity  secreted  in  twenty-four  hours  varies  greatly,  but  is  at  least 
i  liter. 

Function  of  Saliva. — The  purposes  served  by  saliva  are  mechanical 
and  chemical. 

Mechanical. — (i)  It  keeps  the  mouth  in  a  due  condition  of  moisture, 
facilitating  the  movements  of  the  tongue  in  speaking  and  in  the  mastication 
of  food.  (2)  It  serves  also  in  dissolving  sapid  substances,  and  renders  them 
capable  of  exciting  the  nerves  of  taste.  (3)  But  the  principal  mechanical 
purpose  of  the  saliva  is  that,  by  mixing  with  the  food  during  mastication,  it 


FIG.  252. — Showing  the  variation  of  the  rate  of  secretion  of  saliva,  second  line  from 
the  top,  and  variation  of  blood  pressure,  top  line.  At  a,  an  injection  of  o .  2  mgr.  pilocar- 
pine.  At  &,  50  c.c.  oxygenated  blood  was  injected  into  the  jugular  vein.  (Jonescu.) 

makes  a  soft  pulpy  or  creamy  mass  such  as  may  be  easily  swallowed.  To 
this  purpose  the  saliva  is  adapted  both  by  quantity  and  quality.  For,  speak- 
ing generally,  the  quantity  secreted  during  feeding  is  in  direct  proportion 
to  the  dryness  and  hardness  of  the  food. 

Chemical. — The  chemical  action  which  the  saliva  exerts  upon  the  food  in 
the  mouth  is  to  convert  the  starchy  materials  which  it  contains  into  soluble 
starch  and  then  into  sugar.  This  power  the  saliva  owes  to  the  enzyme 
ptyalin.  Certain  investigators  have  of  late  asserted  that  saliva  contains 
another  enzyme,  known  as  maltase,  which  has  the  power  of  splitting  the  di- 
saccharides  into  monosaccharides,  or  maltose  into  dextrose.  The  action  of 
this  ferment  is  certainly  very  limited.  The  conversion  of  the  starch  under 
the  influence  of  the  ferment  into  sugar  takes  place  in  several  stages,  and  in 
order  to  understand  it  a  knowledge  of  the  structure  and  composition  of 
starch  granules  is  necessary.  A  starch  granule  consists  of  two  parts :  an  en- 


356  FOOD   AND    DIGESTION 

velope  of  cellulose,  which  does  not  give  a  blue  color  with  iodine  except  on 
addition  of  sulphuric  acid,  and  of  granulose,  which  is  contained  within,  and 
which  gives  a  blue  color  with  iodine  alone.  Briicke  states  that  a  third  body 
is  contained  in  the  granule,  which  gives  a  red  color  with  iodine,  viz.,  erythro- 
granulose.  The  granulose  swells  up  on  boiling,  bursts  the  envelope,  and  the 
whole  granule  is  more  or  less  completely  converted  into  a  paste  or  gruel 
which  is  called  gelatinous  starch. 

When  ptyalin  acts  upon  boiled  starch,  it  first  changes  the  latter,  by  hydrol- 
ysis, into  soluble  starch,  or  amidulin;  this  is  more  limpid  and  more  like  a  true 
solution,  though  it  still  gives  the  blue  coloration  on  the  addition  of  iodine. 
This  stage  is  very  brief,  only  thirty  seconds  being  sometimes  required  in  labo- 
ratory experiments  to  render  a  stiff  starch  paste  completely  fluid  when  a  few 
drops  of  saliva  are  added  at  body  temperature.  This  rapidity  of  action  is  of 
great  importance,  as  under  proper  conditions  of  mastication  practically  all 
the  boiled  starch  of  the  food  ought  to  enter  the  stomach  as  soluble  starch. 
When  the  starch  has  not  been  previously  boiled,  the  envelope  of  cellulose 
retards  the  action  of  the  ptyalin  to  a  very  marked  degree. 

Starch. 
Soluble  starch. 


Erythro-dextrin.  Maltose  and  iso-maltose. 


Achroo-dextrins.         Maltose  and  iso-maltose. 

The  further  stages  of  hydrolytic  cleavage  result  in  the  formation  of  a 
variable  mixture  of  maltose  and  iso-maltose  with  a  series  of  dextrins,  but  ap- 
parently never  result  (in  laboratory  experiments)  in  the  complete  conversion 
of  the  dextrins  into  sugars.  Gradually,  as  the  starch  is  converted,  the  blue 
coloration  with  iodine  is  replaced  by  a  purplish-red  and  finally  by  a  red 
color:  the  latter  color  is  produced  by  ery thro- dextrin  (so  called  from  the 
color).  In  the  later  stages  no  coloration  is  obtained  with  iodine,  and  for  this 
reason  the  dextrins  formed  are  known  as  achroo- dextrins',  there  are  probably 
several  of  these,  but  they  have  not  yet  been  sufficiently  isolated.  As  sugar 
appears  very  early  in  the  process,  even  at  the  stage  of  erythro-dextrin,  and 
gradually  increases  in  amount,  it  is  generally  concluded  that  maltose  is 
formed  early  in  the  decomposition  of  the  starch  molecule.  The  process  is 
usually  represented  schematically  as  above. 

The  sugars  formed  are  maltose  (C12H22On)  and  a  closely  allied  sugar 
known  as  iso-maltose.  A  small  percentage  of  dextrose  has  been  found  by 
some  observers,  and  this  is  due  to  the  action  of  maltase.  Maltose  is  allied 


ACTION    OF   SAUVA    ON    STARCH  357 

to  saccharose  or  cane-sugar  more  nearly  than  to  glucose;  it  is  crystalline;  its 
solution  has  the  property  of  polarizing  light  to  the  right  to  a  greater  degree 
than  solutions  of  glucose  (3  to  i);  it  is  not  so  sweet,  and  reduces  copper  sul- 
phate less  easily.  It  can  be  converted  into  glucose  by  boiling  with  dilute 
acids  and  by  the  action  of  the  enzyme  maltase  present  in  saliva. 

According  to  Brown  and  Heron,  the  reactions  may  be  represented  thus : 
One  molecule  of  gelatinous  starch  is  converted  by  the  action  of  an  amylolytic 

ferment  into  n  molecules  of  soluble  starch. 

One  molecule  of  soluble  starch  =  (C12H20O10)10 +8H2O,  which  is  further  con- 
verted by  the  ferment  into 

i.   Erythro-dextrin,   (C12H20O10)9  (giving  red  with   iodine)  + 

Maltose  (C12H22On). 
then  into  2.   Erythro-dextrin  (C12H20O10)8  (giving  yellow  with   iodine) 

+  Maltose  2(C12H22OU). 

next  into  3.  Achroo-dextrin  (C12H20O10)7  +  Maltose  3  (C12H22On). 
And  so  on;  the  resultant  being: 

Soluble  starch  (C12H20O10)10  +  Water  8H2O  =  Maltose  8(C12H22On)  + 
Achroo-dextrin  (C12H20O10)2. 

Many  observers,  however,  believe  that  the  maltose  simultaneously  pres- 
ent with  erythro-dextrin  is  not  actually  split  off  from  the  starch  molecule  in 
the  formation  of  erythro-dextrin,  but  that  it  is  the  product  of  more  advanced 
hydrolysis  in  other  starch  molecules.  They  point  out  that  in  such  a  chemical 
reaction  of  considerable  time  duration,  it  is  improbable  that  all  the  starch 
molecules  are  attacked  at  the  same  rate  or  are,  at  any  given  moment,  equally 
advanced  in  cleavage.  Their  theory  is  that  there  is  a  series  of  more  and  more 
simple  dextrins  formed  giving  rise  finally  to  the  disaccharides. 

The  presence  of  sugar  in  such  an  experiment  is  at  once  discovered  by  the 
application  of  Trommer's  test,  which  consists  in  the  addition  of  a  drop  or 
two  of  a  solution  of  copper  sulphate,  followed  by  a  larger  quantity  of  caustic 
potash.  When  the  liquid  is  boiled,  an  orange-red  precipitate  of  copper  sub- 
oxide  indicates  the  presence  of  sugar. 

Influences  which  Affect  the  Action  of  Saliva  on  Starch. — Moderate 
heat,  about  37 .8°  to  40°  C.,  is  most  favorable  to  the  rapid  cleavage  of  starch 
by  the  ptyalin.  Cold  retards  and  o°  C.  suspends  the  action  but  does  not  de- 
stroy the  ferment.  A  temperature  of  60°  C.  destroys  the  ptyalin. 

Removal  of  the  products  of  salivary  digestion  as  they  are  formed  facili- 
tates the  action  of  the  enzyme,  as  an  exeess  of  these  products  is  detrimental 
to  further  action. 

The  reaction  between  starch  and  saliva  takes  place  best  in  a  neutral  or 
very  faintly  alkaline  medium  and  is  inhibited  by  strong  alkalies  and  espe- 
cially by  acids  even  as  weak  as  the  acidity  of  the  gastric  juice.  This  last  is 
of  particular  importance  since  it  raises  the  question  as  to  how  long  the 
ptyalin  may  act. 


358  FOOD   AND    DIGESTION 

The  action  of  saliva  on  starch  is  not  limited  to  the  brief  interval  during 
which  food  remains  in  the  mouth,  as  is  now  well  known,  but  may  continue 
for  a  time  in  the  stomach. 

Ptyalin  is  strictly  an  amylolytic  ferment. 

Starch  appears  to  be  the  only  principle  of  food  with  the  exception  of  the 
dextrins  and  glycogen,  upon  which  the  saliva  acts  chemically.  The  secretion 
has  no  apparent  influence  on  gum,  cellulose,  or  on  fat,  and  is  equally  destitute 
of  power  over  albuminous  and  gelatinous  substances. 

The  salivary  glands  of  children  do  not  produce  functionally  active  saliva 
till  the  age  of  4  to  6  months,  and  hence  the  bad  effects  of  feeding  them  before 
this  age  on  starchy  food,  corn-flour,  etc.,  which  they  are  unable  to  render 
soluble  and  capable  of  absorption. 

Salivary  Digestion  in  the  Stomach. — Laboratory  experiments  have 
demonstrated  that  while  the  addition  of  even  o .  05  per  cent,  of  hydrochloric 
acid  will  inhibit  the  action  of  ptyalin  on  a  solution  of  starch,  if  any  proteins 
be  present  in  the  solution  much  more  acid  must  be  added  before  the  action 
of  the  ptyalin  is  stopped.  The  explanation  of  the  latter  fact  is  that  the  acid 
unites  with  the  proteins  in  some  chemical  combination  forming  "combined 
acid,"  which  has  little  effect,  comparatively,  on  ptyalin.  This  "combined 
acid"  gives  a  red  color  with  litmus,  but  is  distinguished  from  free  acid  by 
giving  a  brownish  instead  of  a  bluish  color  with  Congo  red.  When  food  enters 
an  empty  stomach,  as  happens  at  the  beginning  of  a  meal  the  acid  first  com- 
bines with  the  protein  food  stuffs  and  so  does  not  at  once  affect  the  ptyalin. 

A  still  more  important  fact  in  its  bearing  on  this  subject  was  recently 
discovered  by  Cannon,  who  showed  experimentally  that  starchy  foods  mixed 
with  weak  alkali  remain  alkaline  in  the  stomach  for  as  much  as  an  hour  and 
a  half.  Such  foods  when  swallowed  into  the  stomach  are  packed  away  in 
that  organ  in  a  mass.  The  secretion  of  the  acid  gastric  juice  comes  in  con- 
tact only  with  the  outer  surface  of  the  mass,  which  is  not  materially  disturbed 
by  the  stomach  peristalses.  The  center  of  the  mass  may,  therefore,  remain 
alkaline  until  the  outer  layers  are  completely  eroded  away,  and  the  ptyalin 
may  continue  to  act  on  starch  during  the  whole  time. 

DEGLUTITION. 

When  properly  masticated,  the  food  is  transmitted  in  successive  portions 
to  the  stomach  by  the  act  of  deglutition  or  swallowing.  The  following  account 
of  deglutition  is  based  upon  the  researches  of  Kronecker  and  Meltzer,  whose 
experiments  seem  to  modify  in  some  details  the  earlier  theory  of  Magendie: 

The  mouth  is  closed,  and  the  food  after  thorough  mixing  with  the  saliva 
is  rolled  into  a  bolus  on  the  dorsum  of  the  tongue.  The  tip  of  the  tongue  is 
pressed  upward  and  forward  against  the  hard  palate,  thus  shutting  off  the 
anterior  part  of  the  mouth  cavity.  The  mylo-hyoid  muscles  then  suddenly 


DEGLUTITION  359 

contract,  the  bolus  of  food  is  put  under  great  pressure  and  shot  backward  and 
downward  through  the  pharynx  and  into  the  esophagus  and,  if  the  food  be 
fluid  enough,  even  to  the  cardiac  orifice  of  the  stomach.  Coincidently  with 
the  contraction  of  the  mylo-hyoid  muscles,  the  hyoglossi  are  thrown  into 
action,  drawing  the  tongue  backward  and  downward,  not  only  increasing 
the  pressure  upon  the  food,  but  forcing  the  epiglottis  over  the  glottis,  closing 
the  larynx. 

It  has  been  shown  by  the  Roentgen-ray  method  that  the  character  of  the 
food  determines  somewhat  its  passage  through  the  esophagus.  The  dry 
and  semisolid  foods  are  seized  by  the  musculation  of  the  esophagus  and 
passed  down  that  organ  by  a  peristaltic  wave.  The  longitudinal  muscles 
contract,  tending  to  enlarge  the  diameter  of  the  esophagus  in  advance  of  the 
food,  while  contractions  of  the  circular  muscles  produce  pressure  on  the 
bolus  just  behind,  thus  forcing  it  along  to  the  cardia.  This  wave  reaching 
the  cardiac  orifice  about  six  seconds  after  the  commencement  of  the  act  of 
deglutition,  forces  the  food  into  the  stomach,  the  sphincter  having  previously 
relaxed.  The  interval  of  time  between  the  commencement  of  the  act  of 
deglutition  and  the  arrival  of  the  more  fluid  food  at  the  cardiac  orifice  of  the 
stomach  may  not  be  more  than  one-tenth  second,  though  it  remains  at  the 
cardiac  orifice  without  entering  the  stomach  until  the  first  parts  of  the  act  of 
swallowing  is  reinforced  by  the  subsequent  contraction  of  the  constrictors  of 
the  pharynx  and  the  passage  of  a  peristaltic  wave  down  the  esophagus.  In 
some  cases,  however,  the  liquid  food  is  not  stopped  at  the  cardiac  orifice, 
but  is  sent  through  the  relaxed  sphincter  by  the  original  force  of  the  mylo- 
hyoid  contraction. 

In  man  the  esophagus  was  said  to  contract  in  three  separate  segments, 
the  first  segment  lying  in  the  neck  and  being  about  six  centimeters  long,  the 
second  being  the  next  ten  centimeters  of  the  tube,  and  the  third  the  re- 
maining portion  to  the  stomach.  But  the  later  Roentgen-ray  observations 
show  no  break  in  the  continuous  passage  of  the  food,  though  the  movement 
of  the  food  is  slower  in  the  lower  segment  of  the  esophagus. 

The  act  of  swallowing  consists,  then,  of  the  contraction  in  sequence  of 
the  mylo-hyoids,  the  constrictors  of  the  pharynx,  and  of  the  esophagus.  The 
computed  time  of  contraction  is  as  follows: 

Seconds. 
Contraction  of  mylo-hyoids  and  constrictors  of  the  pharynx ...      0.3 

Contraction  of  the  first  part  of  the  esophagus 0.9 

Contraction  of  the  second  part  of  the  esophagus 1.8 

Contraction  of  the  third  part  of  the  esophagus 3.0 

6.0 

» 

If  a  second  attempt  at  swallowing  be  made  before  the  first  has  been  com- 
pleted (that  is,  before  six  seconds  have  elapsed),  the  remaining  portion  of  the 


360  FOOD    AND    DIGESTION 

first  act  is  inhibited,  and  the  contraction  wave  reaches  the  stomach  six 
seconds  after  the  commencement  of  the  second  act. 

During  the  act  of  deglutition  the  posterior  nares  are  closed  through  the 
action  of  the  levator  palati  and  tensor  palati  muscles,  which  raise  the  velum; 
the  palato-pharyngei,  drawing  the  posterior  pillars  of  the  fauces  together; 
and  the  azygos  uvulae,  which  raises  the  uvula — thus  forming  a  complete  cur- 
tain. Otherwise  the  food  would  pass  into  the  nose,  as  happens  in  the  case 
of  cleft  palate.  At  the  same  time  the  larynx  is  closed  by  the  adductor 
muscles  of  the  vocal  cords  and  the  descent  of  the  epiglottis,  the  larynx  being 
drawn  upward  as  a  whole  through  the  action  of  the  mylo-hyoid,  genio-hyoid, 
thyro-hyoid,  and  digastric  muscles.  The  presence  of  the  epiglottis  is  not 
necessary  for  the  completion  of  the  act  of  deglutition. 

Nervous  Mechanism  of  Deglutition. — The  sensory  nerves  engaged 
in  the  reflex  act  of  deglutition  are  branches  of  the  fifth  cerebral,  supplying  the 
soft  palate;  the  glosso-pharyngeal,  supplying  the  tongue  and  pharynx;  the 
superior  laryngeal  branch  of  the  vagus,  supplying  the  epiglottis  and  the  glottis. 
The  motor  fibers  concerned  are  branches  of  the  fifth,  supplying  part  of  the 
digastric  and  mylo-hyoid  muscles  and  the  muscles  of  mastication;  the  facial, 
supplying  the  levator  palati;  the  glosso-pharyngeal,  supplying  the  muscles 
of  the  pharynx;  the  vagus,  supplying  the  muscles  of  the  larynx  through  the 
inferior  laryngeal  branch;  and  the  hypoglossal,  the  muscles  of  the  tongue. 
The  nerve  center  by  which  the  muscles  are  harmonized  in  their  action  is 
situated  in  the  medulla  oblongata.  It  cannot  be  definitely  circumscribed, 
but  is  in  the  general  level  of  the  vagus  origin.  The  movements  of  the  esoph- 
agus are  co-ordinated  by  the  complex  of  sensory  and  motor  fibers  of  the 
fifth  and  the  ninth  to  twelfth  cranial  nerves,  which  all  take  some  part  in  this 
complicated  reflex. 

Cannon  has  demonstrated  that  the  smooth  muscle  of  the  lower  end  of 
the  esophagus  and  around  the  cardiac  orifice  is  maintained  in  contraction 
by  a  local  reflex  mechanism.  This  prevents  regurgitation  of  the  foods. 
The  local  apparatus  is  brought  into  action  by  the  stimulation  of  sensory  cells 
in  the  mucous  membrane  of  this  region  of  the  stomach  by  the  acid  of  the 
gastric  secretion.  The  reflex  is  assumed  to  be  a  local  one  taking  place  through 
the  intrinsic  nervous  mechanism.  This  acid  closure  of  the  cardiac  sphincter 
is  to  be  compared  with  the  similar  mechanism  for  the  pylorus,  see  page  356. 

DIGESTION  IN  THE  STOMACH. 

The  stomach  of  man  and  the  carnivora  is  the  dilated  portion  of  the  ali- 
mentary canal  following  the  esophagus.  The  esophagus  enters  the  stomach 
at  the  cardiac  end  and  the  pyloric  end  of  the  stomach  is  continuous  with  the 
duodenal  part  of  the  intestine.  It  varies  in  shape  and  size  according  to  its 
state  of  distention.  It  is  supplied  with  nerves  from  the  vagus  and  from  the 
sympathetic  and  receives  a  special  artery,  the  gastric  artery. 


STRUCTURE  OF  THE  STOMACH 


361 


Structure  of  the  Stomach. — The  stomach  is  composed  of  four  coats, 
called,  respectively,  the  external  or  peritoneal,  the  muscular,  the  submucous> 


FIG.  253. — The  Human  Stomach  and  the  Vagus  Distribution.  R.  L.,  Recurrent 
laryngeal;  Ca2,  inferior  cervical  cardiac  branch;  €0,3,  €0,4,  cardiac  branches  of  vagus; 
A.  P.  PI.,  P.  P.  PI.,  anterior  and  posterior  pulmonary  plexuses;  Oes.  PL,  esophageal  plexus; 
Cast.  R.  and!,.,  gastric  branches  of  vagus,  right  and  left;  Coe,  PL,  coeliac  plexus;  Hep.  pi., 
hepatic  plexus. 

and  the  mucous  coat.     Blood  vessels,  lymphatics,  and  nerves  are  distributed 
in  and  between  them. 


362 


FOOD   AND    DIGESTION 


The  muscular  coat  consists  of  three  separate  layers  of  fibers  which,  accord- 
ing to  their  several  directions,  are  named  the  longitudinal,  circular,  and 
oblique.  The  longitudinal  set  are  the  most  superficial  and  are  continuous 
with  the  longitudinal  fibers  of  the  esophagus  and  spread  out  in  a  diverging 

manner  over  the  cardiac  end  and  sides  of 
the  stomach  to  the  pylorus.  The  circular 
or  transverse  coat  more  or  less  completely 
encircles  all  parts  of  the  stomach;  this 
coat  is  thickest  at  the  middle  and  in  the 
pyloric  portion  of  the  organ,  and  forms 
the  chief  part  of  the  thick  ring  of  the 
pylorus.  The  next  and  consequently 
deepest  coat,  the  oblique,  is  continuous 
with  the  circular  muscular  fibers  of  the 


FIG.  254. 


FIG.  255. 


FIG.  254. — From  a  Vertical  Section  through  the  Mucous  Membrane  of  the  Cardiac  End 
of  Stomach.  Two  peptic  glands  are  shown  with  a  duct  common  to  both,  one  gland  only  in 
part,  a,  Duct  with  columnar  epithelium  becoming  shorter  as  the  cells  are  traced  down- 
ward; n,  neck  of  gland  tubes,  with  central  and  parietal  or  so-called  peptic  cells;  h,  fundus 
with  curved  cecal  extremity — the  parietal  cells  are  not  so  numerous  here.  X  400.  (Klein 
and  Noble  Smith.) 

FIG.  255. — Cross- sections  at  Various  Levels  of  Peptic  Glands  of  Stomach.  X  400. 
M,  Section  through  gastric  pit  near  surface;  M' ,  section  through  gastric  pit  near  bottom; 
h,  mouth  of  gland;  k,  neck;  g,  body  near  fundus;  the  chief  cells  are  shaded  lightly;  b,  parietal 
cells.  (Kolliker.) 

esophagus  at  the  cardiac  orifice  of  the  stomach.  This  coat  is  quite  inter- 
rupted and  more  or  less  incomplete.  The  muscular  fibers  of  the  stomach 
and  intestinal  canal  are  unstriated. 


THE    GASTRIC    GLANDS 


363 


The  mucous  membrane  of  the  stomach,  which  rests  upon  a  layer  of  loose 
cellular  membrane,  or  submucous  tissue,  is  smooth,  soft  and  velvety.  It  is 
of  a  pale  pink  color  during  life,  and  in  the  contracted  state  is  thrown  into 
numerous  longitudinal  folds  or  rugae,  which  disappear  when  the  organ  is 
distended.  It  is  composed  of  a  mass  of  short  tubular  secreting  glands. 

The  Gastric  Glands. — The  glands  of  the  mucous  membrane  of  the 
stomach  are  of  two  varieties,  Cardiac  and  Pyloric. 


FIG.  256. 


FIG.  257. 


FIG.  256. — Longitudinal  Section  of  Fundus  of  Gland  from  Dog's  Stomach,  a, 
Lumen  of  gland;  b,  intracellular  canals  in  parietal  cells;  c,  cut-off  portion  of  parietal  cell; 
d,  chief  cells;  e,  intercellular  canals  leading  from  lumen  of  gland  to  canals  in  parietal  cells. 
(Bailey.) 

FIG.  257. — Tubule  of  Pyloric  Gland  of  Man.  Note  the  thin  basal  layer  of  cytoplasm; 
the  reticular  cell  body  containing  secretion;  the  subdivision  of  the  latter  in  some  cells  into 
proximal  and  distal  masses.  Highly  magnified.  (Bailey.) 


Cardiac  glands  are  found  throughout  the  whole  of  the  cardiac  end  of  the 
stomach.  They  are  arranged  in  groups  of  four  or  five,  which  are  separated 
by  a  fine  connective  tissue.  Two  or  three  tubes  often  open  into  one  duct, 
figure  254,  which  forms  about  a  third  of  the  whole  length  of  the  tube  and 
opens  on  the  surface.  The  ducts  and  the  free  surface  are  lined  with  colum- 
nar epithelium.  The  body  of  the  gland  is  composed  of  granular  secreting 
cells,  called  chief  cells  or  peptic  cells.  Between  these  cells  and  the  membrana 
propria  of  the  tubes  are  large  oval  or  spherical  cells,  granular  in  appearance 
with  clear  oval  nuclei;  these  cells  are  called  parietal  cells.  They  do  not 
form  a  continuous  layer,  figure  254.  Intercellular  tubules  extending  from 


364 


FOOD   AND    DIGESTION 


the  duct  of  the  gland  between  the  chief  cells  and  connecting  with  intracellular 
secretory  tubules  in  the  parietal  cells  have  been  shown  by  the  Golgi  silver 
method,  by  napthol  blue,  etc.,  figure  256. 

As  the  pylorus  is  approached  the  gland  ducts  become  longer  and  the 
tube  proper  becomes  shorter,  and  occasionally  branched  at  the  fundus. 

The  Pyloric  Glands. — These  glands  have  much  longer  ducts  and  larger 
mouths  than  the  peptic  glands. 

The  parietal  cells  are  absent  in  the  pyloric  glands.  The  pyloric  glands 
become  larger  as  they  approach  the  duodenum,  also  more  convoluted  and 
more  deeply  situated.  They  are  directly  continuous  with  Brunner's  glands 
in  the  duodenum  (Watney). 


FIG.  258. — Scheme  of  Blood  Vessels  and  Lymphatics  of  Stomach.  X  70.  a,  Mucous 
membrane;  &,  muscularis  mucosae;  c,  submucosa;  d,  inner  circular  muscle  layer;  e,  outer 
longitudinal  muscle  layer;  A,  blood  vessels;  B,  structure  of  coats;  C,  lymphatics.  (Szymo- 
nowicz,  after  Mall.) 

Blood  vessels  and  Lymphatics. — The  blood  vessels  of  the  stomach  first 
break  up  in  the  submucous  tissue  and  send  branches  upward  between  the 
closely  packed  glandular  tubes,  which  anastomose  around  them  by  a  fine 
capillary  network  with  oblong  meshes.  Contiguous  with  this  deeper  plexus, 
or  prolonged  upward  from  it,  so  to  speak,  is  a  more  superficial  network  of 
larger  capillaries,  which  branch  densely  around  the  orifices  of  the  tubes  and 
form  the  framework  on  which  are  molded  the  small  elevated  ridges  of  mucous 


ACT    OF    SECRETION    OF    GASTRIC   JUICE  365 

membrane.  From  this  superficial  network  the  veins  chiefly  take  their  origin, 
pass  down  between  the  tubes,  with  no  very  free  connection  with  the  deeper 
intertubular  capillary  plexus,  and  open  finally  into  the  venous  network  in 
the  submucous  tissue. 

The  lymphatic  vessels  surround  the  gland  tubes  with  a  network. 
Toward  the  fundus  of  the  peptic  glands  are  masses  of  lymphoid  tissue 
which  may  appear  as  distinct  follicles,  somewhat  like  the  solitary  glands 
of  the  small  intestine. 

Microscopic  Changes  in  the  Gastric  Glands  During  Secretion.— 
Langley  has  made  a  study  of  the  histological  changes  in  the  glandular  tissues 
in  the  fresh  state.  He  finds  that  during  fasting  or  when  the  glands  are  at  rest 
the  chief  cells  are  granular  throughout,  being  crowded  with  large  highly  re- 
fractive granules.  During  activity  these  granules  gradually  disappear  pro- 
gressively from  the  base  toward  the  border  of  the  cell  on  the  lumen  of  the  tube. 
They  no  doubt  represent  the  zymogen  substances  from  which  the  first  dis- 
charge of  enzyme  is  derived  during  the  activity  of  secretion.  The  parietal 
cells  are  finely  granular  throughout,  though  they  decrease  in  size  during 
activity,  as  in  fact  do  the  chief  cells.  Macallum  by  the  use  of  microchemical 
tests  has  shown  the  presence  of  excess  of  chlorides  in  the  ducts  and  in- 
tracellular  canals,  and  in  the  parietal  cells.  The  pyloric  cells  do  not  undergo 
such  marked  changes,  and  the  mucous  cells  of  the  more  superficial  layers 
of  the  mucosa  cannot  be  said  to  show  any  special  changes  at  the  time  of 
digestional  activity  of  the  other  layers.  During  periods  of  rest  the  gastric 
cells  increase  in  size  and  again  become  charged  with  granules  as  before. 

The  Act  of  Secretion  of  Gastric  Juice. — The  gastric  glands  un- 
dergo periods  of  rest  and  activity.  The  active  secretion  of  normal  gastric 
juice  takes  place  when  food  is  introduced  into  the  mouth,  or  in  fact  the 
mere  sight  of  appetizing  food  is  followed  by  an  abundant  secretion  of  gastric 
juice,  as  shown  by  Bidder  and  Schmidt  on  the  dog  with  a  gastric  fistula.  Such 
observations  strongly  indicate  that  the  act  is  a  nervous  phenomenon,  at  least 
under  nervous  control. 

Quite  recently  Pavlov  has  proved  that  secretory  fibers  are  carried  to 
the  gastric  glands  in  the  vagus  trunk.  His  experiment  consisted  in  estab- 
lishing a  gastric  fistula,  and  some  days  later  in  dividing  the  esophagus 
in  the  neck  in  such  a  manner  that  any  food  swallowed  would  be  diverted 
to  the  exterior  through  the  cut  end.  A  "  fictitious  meal"  could  then  be  given 
to  the  animal,  and  the  effect  upon  the  stomach  noted.  As  long  as  the  vagi 
were  intact,  certain  foods  (meats)  caused  a  flow  of  gastric  juice,  though 
none  of  the  food  reached  the  stomach.  The  secretion  of  gastric  juice  con- 
tinued for  hours  with  the  production  of  a  large  quantity  of  secretion.  When 
the  vagi  had  been  cut,  no  secretion  occurred.  Moreover,  he  found  that  direct 
stimulation  of  the  vagus  produced  a  flow  of  gastric  juice. 

Khigine  placed  foods  in  an  isolated  gastric  pouch  prepared  with  care  to 


366 


FOOD   AND   DIGESTION 


maintain  the  nervous  relations  intact,  and  it  led  to  secretion  of  gastric  juice 
in  the  main  part  of  the  stomach.  This  is  undoubtedly  a  nervous  reflex  effect. 
Recently  observations  on  a  case  of  stricture  of  the  human  esophagus 
which  prevented  food  from  reaching  the  stomach  have  shown  that  an 
abundant  flow  of  gastric  juice  takes  place  when  food  is  taken  into  the  mouth. 


RV 


FIG.  259. — Very  Diagrammatic  Representation  of  the  Nerves  of  the  Alimentary 
Canal.  Oe  to  Ret,  the  various  parts  of  the  alimentary  canal  from  esophagus  to  rectum; 
L.  V,  left  vagus,  ending  on  front  of  stomach;  rl,  recurrent  laryngeal  nerve,  supplying  upper 
part  of  esophagus;  R.  V,  right  vagus,  joining  left  vagus  in  esophageal  plexus;  as.  pi., 
supplying  the  posterior  part  of  stomach,  and  continues  as  R'  V  to  join  the  solar  plexus,  here 
represented  by  a  single  ganglion,  and  connected  with  the  inferior  mesenteric  ganglion,  m. 
gl.;  a,  branches  from  the  solar  plexus  to  stomach  and  small  intestine,  and  from  the  mesen- 
teric ganglia  to  the  large  intestine;  Spl.  maj.,  large  splanchnic  nerve,  arising  from  the 
thoracic  ganglia  and  rami  communicantes;  r.  c.,  belonging  to  dorsal  nerves  from  the  6th 
to  the  Qth  (or  loth);  Spl.  min.,  small  splanchnic  nerve  similarly  from  the  loth  and  nth 
dorsal  nerves.  These  both  join  the  solar  plexus,  and  thence  make  their  way  to  the  ali- 
mentary canal;  c.  r.,  nerves  from  the  ganglia,  etc.,  belonging  to  nth  and  i2th  dorsal  and 
ist  and  2d  lumbar  nerves,  proceeding  to  the  inferior  mesenteric  ganglia  (or  plexus),  m.  gl., 
and  thence  by  the  hypogastric  nerve,  n.  hyp.,  and  the  hypogastric  nerve,  n.  hyp.,  and  the 
hypogastric  plexus,  pi.  hyp.,  to  the  circular  muscles  of  the  rectum;  /.  r.,  nerves  from  the  2d 
and  3d  sacral  nerves,  S.  2,  S.  3  (nervi  erigentes)  proceeding  by  the  hypogastric  plexus  to  the 
longitudinal  muscles  of  the  rectum.  (M.  Foster.) 

It  seems  conclusively  established  at  the  present  time  that  the  secretion  of 
gastric  juice  is  a  reflex  act  controlled  by  a  definite  nervous  mechanism.  This 
reflex  can  be  aroused  by  the  sensory  stimuli  of  taste,  smell,  and  even  sight. 
It  can  also  be  initiated  by  stimuli  arising  in  the  stomach  itself  by  the  effects 
of  ingredients  of  the  food  or  by  the  products  of  digestion.  Indeed,  it  has 
been  shown  that  peptone  is  a  very  efficient  stimulus  for  this  stomach  reflex. 

Edkins,  however,  has  recently  shown  that  the  contact  of  certain  food 
products  with  the  pyloric  end  of  the  stomach,  where  they  are  slightly  ab- 
sorbed, gives  rise  to  some  chemical  substance — a  gastric  hormone  or  secre- 


THE    GASTRIC   JUICE 


367 


tagogue — which  acts  as  a  powerful  stimulus  to  gastric  secretion  when  it  is 
introduced  into  the  circulation.  Such  food  substances  are  dextrins,  maltose 
and  dextrose,  proteoses,  and  above  all  meat  extract. 

The  influence  of  the  higher  nerve  centers  on  gastric  digestion,  as  in  the 
case  of  emotions,  is  too  well  known  to  need  more  than  a  reference. 


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FIG.  260. — Table  to  show  the  Secretion  of  Gastric  Juice  by  the  Dog.     (Lliffine.) 

Immediately  on  the  introduction  of  food  or  other  stimulating  substance, 
the  mucous  membrane,  which  was  previously  quite  pale,  becomes  slightly 
turgid  and  reddened  with  the  influx  of  a  larger  quantity  of  blood,  and  the  gas- 
tric glands  commence  actively  to  secrete.  An  acid  fluid  is  poured  out  in 
minute  drops  and  the  secretion  may  continue  for  hours. 


368  FOOD    AND    DIGESTION 

The  Gastric  Juice. — The  first  analysis  of  gastric  juice  was  made  by 
Prout  on  a  small  and  impure  specimen.  Beaumont  made  an  elaborate  and 
classic  series  of  observations  on  the  gastric  secretion  of  Alexis  St.  Martin, 
in  whom  there  existed,  as  the  result  of  a  gunshot  wound,  an  opening  leading 
directly  into  the  stomach  near  the  upper  extremity  of  the  great  curvature 
and  three  inches  from  the  cardiac  orifice.  The  introduction  of  any  mechan- 
ical irritant,  such  as  the  bulb  of  a  thermometer,  into  the  stomach  through 
this  artificial  opening  excited  the  secretion  of  gastric  fluid.  This  was  drawn 
off,  and  was  often  obtained  to  the  extent  of  nearly  an  ounce. 

The  chemical  composition  of  human  gastric  juice  has  been  also  investi- 
gated by  Schmidt.  The  fluid  in  this  case  also  was  obtained  by  means  of  an 
accidental  gastric  fistula.  The  mucous  membrane  was  excited  to  action  by 
the  introduction  of  some  hard  matter,  such  as  dry  peas,  and  the  secretion  was 
removed  by  means  of  an  elastic  tube.  The  fluid  obtained  was  found  to  be 
acid,  limpid,  odorless,  with  a  specific  gravity  of  i .  002  to  i .  oio.  It  contained 
a  few  cells  and  some  fine  granular  matter.  The  analysis  of  the  fluid  obtained 
in  this  way  is  given  below.  Essentially  it  is  a  weakly  acid  fluid  containing 
hydrochloric  acid  and  enzymes,  of  which  pepsin  and  rennin  are  the  chief, 
though  lipase  and  maltase  are  both  present.  The  gastric  juice  obtained 
from  gastric  fistulas  of  dogs  and  other  animals  shows  some  difference  in 
composition. 

CHEMICAL  COMPOSITION  OF  GASTRIC  JUICE     (SCHMIDT). 

Dogs.  Human. 

Water 971-1?  994-4 

Solids 28.82  5.60 

Solids- 
Ferment — pepsin *  7  •  5  3  •  J9 

Hydrochloric  acid  (free) 2.7  0.2 

Salts- 
Calcium,  sodium,  and  potassium   chlorides;   and 

calcium,  magnesium,  and  iron  phosphates 8.57  2.19 

The  quantity  of  gastric  juice  secreted  daily  has  been  variously  estimated; 
but  the  average  for  a  healthy  adult  may  be  assumed  to  range  from  2,000  to 
3,000  cubic  centimeters  in  the  twenty-four  hours. 

The  Nature  and  Origin  of  the  Acid  of  Gastric  Juice. — The  acidity 
of  the  fluid  is  due  to  free  hydrochloric  acid,  although  other  acids,  e.g.,  lactic, 
acetic,  butyric,  are  not  infrequently  to  be  found  therein  as  products  of  gastric 
digestion  or  abnormal  fermentation.  In  healthy  gastric  juice  the  amount  of 
free  hydrochloric  acid  is  usually  about  0.2  per  cent.,  but  may  be  as  much  as 
0.3  per  cent.  In  pathological  conditions  it  may  be  entirely  absent,  or  may 
amount  to  0.5  per  cent.,  or  even  more. 

Hydrochloric  acid  is  the  proper  acid  of  healthy  gastric  juice,  and  various 
tests  have  been  used  to  prove  this.  The  tests  depend  upon  changes  produced 


THE   ACID    OF    GASTRIC   JUICE  369 

in  aniline  colors  by  the  action  of  hydrochloric  acid  even  in  minute  traces, 
whereas  lactic  and  other  organic  acids  have  no  such  action.  An  aqueous 
solution  of  oo-tropeolin,  a  bright  yellow  dye,  is  turned  red  on  the  addition  of 
a  minute  trace  of  hydrochloric  acid,  and  aqueous  solutions  of  methyl  violet 
and  gentian  violet  are  turned  blue  under  the  same  circumstances. 

The  protein  matter  in  the  food  combines  to  some  extent  with  the  hydro- 
chloric acid,  which  then  is  known  as  combined  acid  and  does  not  redden  litmus- 
paper.  As  this  combination  is  immediate,  it  follows  that  no  free  acid  is  found 
in  the  gastric  contents  until  the  amount  secreted  is  more  than  enough  to  satu- 
rate the  various  albuminous  affinities.  It  is  partly  for  this  reason  that,  as  al- 
ready mentioned,  salivary  digestion  may  continue  in  the  stomach  for  some 
time  after  the  commencement  of  gastric  digestion.  According  to  Ehrlich,  the 
amount  necessary  to  saturate  the  affinities  of  100  grams  of  various  articles 
of  diet  is  as  follows: 

Beef  (boiled) 2.0  grams  of  pure  HC1. 

Mutton  (boiled) 1.9  grams  of  pure  HC1. 

Veal  (boiled) 2.2  grains  of  pure  HC1. 

Pork  (boiled) 1.6  grams  of  pure  HC1. 

Ham  (boiled) 1.8  grams  of  pure  HC1. 

Sweetbread  (boiled) 0.9  gram    of  pure  HC1. 

Wheat  bread 0.3  gram    of  pure  HC1. 

Rye  bread 0.5  gram    of  pure  HC1. 

Swiss  cheese 2.6  grams  of  pure  HC1. 

Milk  (100  c.c.) o  .32-0 .42  gram    of  pure  HC1. 

The  acid  of  the  gastric  juice  is  not  found  until  after  the  secretion  is  poured 
out  on  the  surface  of  the  mucous  membrane  of  the  stomach.  Thus  Claude 
Bernard  after  microscopic  examination  said  that  there  was  no  acid  in  the 
gastric  glands,  that  "the  acid  of  the  gastric  juice  is  formed  only  after  the 
secretion  of  the  juice,  the  glands  secreting  a  liquid  which  breaks  up  into  an 
acid  fluid  and  another  product  as  yet  not  definitely  determined."  Harvey 
and  Bensley,  from  whom  the  translation  just  given  is  quoted,  confirm  Ber- 
nard's views  completely.  They  find  by  an  exhaustive  study  and  by  ingenious 
staining  methods  for  identifying  alkalinity  and  acidity,  that  the  acid  of  gastric 
juice  does  not  make  its  appearance  until  the  secretion  reaches  the  open 
mouths  of  the  glands  and  the  surface  of  the  mucosa.  They  observe  that 
in  the  gland  ducts  the  secretion  is  viscid,  adherent,  stainable,  and  "breaks  up 
into  round  droplets  which  maintain  their  individuality  for  several  minutes," 
noting  "the  red  reaction  also  at  the  same  time  slowly  changing  to  the  blue 
acid  reaction,  if  the  secretion  has  been  stained  with  cyanamin."  "From  these 
observations  we  are  obliged  to  conclude  that  the  secretion  formed  in  the  gland 
possesses  a  relatively  high  content  of  solids,  and  that  the  bulk  of  the  water 
found  in  the  gastric  secretion  is  added  at  the  level  of  the  glandular  foveolae." 
The  parietal  cells  are  alkaline  in  reaction  and  not  acid,  as  are  in  fact  all  the 
tissues  of  the  gland.  However,  the  observation  is  well  established  that  the 


370 


FOOD   AND   DIGESTION 


parietal  cells  are  peculiarly  rich  in  chlorides,  and  these  chlorides  enter  into 
the  composition  of  the  secretion  and  apparently  are  the  final  source  of  the 
hydrochloric  acid  formed  in  the  secretion. 

Malay  holds  that  the  acid  probably  results  from  a  combination  of  common 
salt  with  monosodic  phosphate,  NaH2PO4  +  NaCl  =  Na2HPO4  +  HC1; 
the  disodic  phosphate  is  then  reconverted  by  the  action  of  carbonic  acid 
and  water,  Na2HPO4  +  CO2  +  H2O  =  NaH2PO4  +  NaHCO3.  All  these 
salts  are  found  in  the  gastric  secretion.  However,  Harvey  and  Bensley 
believe  that  the  hydrochloric  acid  is  derived  from  an  organic  combination  of 
the  chlorides  in  the  secretion,  the  nature  of  which  is  not  determined. 

The  Pepsin. — The  pepsin  of  the  gastric  juice  is  derived  from  the  ac- 
tivity of  the  chief  cells  of  the  fundic  glands.  The  zymogen,  pepsinogen, 
which  is  its  immediate  precursor,  is  in  all  probability  represented  by  the  gran- 
ules of  the  resting  cells.  The  ferment  pepsin  does  not  exist  as  such  in  the 
cells,  for  an  extract  of  peptic  glands  in  o .  2  per  cent,  soda  solution  kept  at 
40°  C.  retains  for  hours  its  power  to  digest  protein  when  added  to  o .  2  per 
cent,  hydrochloric  acid.  If  the  extract  be  first  treated  with  acid  till  it  is 
active,  then  neutralized  and  kept,  it  quickly  loses  its  power  to  digest.  The 
enzyme  is  destroyed  by  the  treatment,  but  the  pro-enzyme  is  not  so  injured. 

Digestive  Action  of  Pepsin  and  Hydrochloric  Acid. — The  chief  func- 
tion of  gastric  juice  is  to  alter  the  protein  food  stuffs  so  that  they  may  be 
readily  absorbed.  Less  important  functions  are  the  antiseptic  action  of 
the  hydrochloric  acid  and  the  coagulation  of  milk.  The  chief  digestive 
power  of  the  gastric  juice  depends  on  the  pepsin  and  acid  contained  in  it, 
both  of  which  are  necessary  for  the  process  in  the  stomach. 

This  action  on  proteins  may  be  shown  by  adding  a  little  gastric  juice 
(natural  or  artificial)  to  some  flakes  of  fibrin  or  to  diluted  egg  albumin,  and 
keeping  the  mixture  at  a  temperature  of  about  37 . 8°  C.  (100°  F.).  It  is  soon 
found  that  the  fibrin  goes  into  solution  and  that  the  albumin  cannot  be  pre- 
cipitated on  boiling.  If  the  solution  be  neutralized  with  an  alkali,  a  precipi- 
tate of  acid  metaprotein  is  thrown  down.  After  a  while  the  acid  metaprotein 
disappears,  so  that  no  precipitate  results  on  neutralization,  and  proper 
analysis  will  show  that  all  the  fibrin  or  albumin  has  been  converted  into  other 
protein  ubstances,  viz.,  proteases  and  peptones.  The  process,  as  in  the  case 
of  salivary  digestion,  is  never  complete  and  the  final  result  is  always  a  mixture 
of  peptones  with  proteoses  which  cannot  be  further  peptonized.  The  re- 
lative proportions,  of  course,  depend  on  the  duration  of  the  process.  A  side 
product  is  found  (as  an  insoluble  residue)  in  artificial  gastric  digestion  which 
gives  practically  all  the  protein  reactions  and  is  soluble  in  dilute  alkali, 
though  insoluble  in  water,  sodium  chloride,  or  dilute  acid.  This  is  known 
as  anti-albumid  and  may  be  changed  into  peptone  by  prolonged  digestion;  it 
does  not  occur  in  physiological  gastric  digestion.  The  commonest  proteose 
is  the  one  formed  from  albumin  and  is  known  as  albumose,  or  by  the  more 


PRODUCTS    OF    GASTRIC   DIGESTION  371 

general  name  protease;  this  name  is  used  in  the  subsequent  descriptions  of 
the  digestive  processes. 

All  classes  of  proteins  are  digested  by  gastric  juice,  leading  to  the  produc- 
tion of  proteoses  and  peptones.  The  change  is  indicated  best  by  the  charac- 
ters of  the  new  protein  formed.  Peptones  have  certain  characteristics  which 
distinguish  them  from  other  proteins.  They  are  diffusible;  i.e.,  they  possess 
the  property  of  passing  through  animal  membranes.  In  their  diffusibility 
peptones  differ  remarkably  from  egg  albumin,  and  on  this  diffusibility  depends 
one  of  their  chief  uses.  Egg  albumin  as  such,  even  in  a  state  of  solution, 
would  be  of  little  service  as  food,  inasmuch  as  its  diffusibility  renders  difficult 
its  absorption  or  in  the  case  of  insoluble  proteins  effectually  prevents  absorp- 
tion into  the  blood  vessels  of  the  digestive  canal.  When  completely  changed 
by  the  action  of  the  gastric  juice  into  peptones,  albuminous  matters  diffuse 
readily,  and  can  be  then  absorbed.  Peptones,  however,  are  not  found  in 
the  blood,  even  of  the  vessels  immediately  concerned  in  absorption  from 
the  stomach  and  intestines.  As  will  be  shown,  the  proteins  are  broken 
down  into  their  simpler  cleavage  products  ^in  the  intestine. 

Products  at  Different  Stages  of  Gastric  Digestion. — The  protein 
is  first  changed  into  syntonin,  or  acid  metaprotein,  by  the  combined  action 
of  the  pepsin  and  acid.  Though  the  acid  alone  is  capable  of  accomplishing 
this  step,  the  fact  that  it  does  not  do  so  physiologically  is  proven  by  the 
great  length  of  time  required  in  laboratory  experiments  for  the  change. 
The  acid  is  absolutely  essential  to  the  action  of  pepsin. 

The  next  change  is  the  conversion  of  the  syntonin  into  proteoses  which, 
according  to  Neumeister,  occurs  in  two  successive  stages.  The  first  of  these 
stages  is  the  conversion  of  syntonin  into  the  primary  proteoses;  i.  e.,  proto- 
proteose  and  hetero-proteose.  The  second  is  the  conversion  of  both  proto- 
proteose  and  hetero-proteose  into  the  secondary  proteoses;  i.e.,  deutero- 
proteose.  The  last  change  is  the  conversion  of  the  deutero-proteose  into  the 
end  product  peptone.  This  last  change  does  not  occur  to  any  great  extent 
and  the  proteoses  always  predominate  in  the  digesting  mass.  The  action 
of  pepsin  is  one  of  hydrolysis  and  the  products  are  hydrated  forms  of  protein. 
Schematically  the  changes  in  the  proteins  may  be  represented  as  follows: 

Protein 
Acid  meta-protein. 


Proto-proteose.  Hetero-protecse. 

Deutero-proteose.  Deutero-proteose. 

Peptone.  Peptone. 


372  FOOD   AND    DIGESTION 

Circumstances  Influencing  Gastric  Digestion.— A  temperature  of 
about  40°  C.  is  most  favorable  to  gastric  digestion.  The  pepsin  is  destroyed 
by  a  temperature  of  55°  (neutral)  to  65°  C.  (acid  solution)  and  its  action  is 
retarded  and  suspended  by  low  temperatures.  It  is  inactive  in  neutral 
or  alkaline  solution,  for  an  acid  medium  is  necessary.  Hydrochloric  is  the 
best  acid  for  the  purpose,  but  other  mineral  acids  or  the  organic  acids  may 
be  substituted  for  the  hydrochloric.  Excess  of  peptone  delays  the  action,  and 
the  removal  of  the  products  of  digestion  facilitates  the  process. 

Action  of  Rennin. — Milk  is  curdled  by  the  action  of  gastric  juice,  the 
casein  being  first  precipitated,  and  then  dissolved.  The  curdling  is  due  to  a 
special  ferment  of  the  gastric  juice,  rennin,  and  is  not  due  to  the  action  of  the 
free  acid  alone.  The  effect  of  rennin,  which  is  obtained  commercially 
from  the  fourth  stomach  of  the  calf,  has  long  been  known,  as  it  is  used  ex- 
tensively to  cause  precipitation  of  casein  in  cheese  manufacture.  The  fer- 
ment rennin  is  active  in  a  neutral  solution  as  well  as  in  acid. 

The  Action  of  Gastric  Lipase. — For  many  years  it  has  been  known 
that  fats  were  digested  in  the  stomach,  but  it  has  been  a  more  difficult  matter 
to  definitely  prove  the  source  of  the  lipase,  most  physiologists  holding  that 
the  lipase  is  regurgitated  from  the  intestine.  In  1880  Cash  proved  that 
extracts  of  the  gastric  mucosa  contained  an  active  lipase  which  experimentally 
caused  the  dissociation  of  neutral  fats  as  tested  by  the  increased  amount 
of  fatty  acid.  He  removed  the  pancreas  and  showed  that  fats  were  still 
digested.  Ogata  made  the  tests  in  the  living  stomach  of  the  dog,  closing  off 
the  opening  into  the  intestine.  The  stomach  thus  isolated  and  washed  out 
with  physiological  saline  repeatedly  caused  the  appearance  of  fatty  acid 
when  olein  was  introduced  and  brought  into  contact  with  the  living  gastric 
mucosa.  It  is  evident  that  the  secretion  of  the  gastric  glands  contains  an 
active  lipase. 

The  well  known  observation  of  Pawlow  showing  that  the  secretion  of 
pepsin  is  inhibited  by  an  excess  of  fat  in  the  stomach,  when  taken  in  con- 
nection with  other  facts  showing  the  specific  nature  of  the  digestive 
secretions,  see  figure  268,  suggests  that  lipase  secretion  may  be  thus 
stimulated. 

Pancreatic  Digestion  in  the  Stomach. — Boldyroff  has  recently  shown 
that  after  the  ingestion  of  fats  or  fatty  foods  in  sufficient  amounts,  that  the 
secretion  of  gastric  juice  is  inhibited  and  that  the  presence  of  the  pancreatic 
and  intestinal  secretions  can  be  demonstrated  in  the  stomach  contents. 

The  accuracy  of  this  observation  as  suggested  from  the  preceding  para- 
graph may  be  held  in  question  to  some  extent,  since  the  fat  enzymes  are 
present  in  the  gastric  juice  itself.  But  that  there  may  be  regurgitation  of 
the  intestinal  juices  into  the  stomach  is  further  supported  by  numerous 
clinical  observations.  This  regurgitation  may,  often  does,  take  place  in 
great  amount  in  the  later  stages  of  gastric  digestion,  at  the  time  when  the 


MOVEMENTS    OF   THE    STOMACH  373 

pyloric  valve  is  least  vigorously  active.  The  presence  of  bile  under  these 
conditions  is  usually  taken  as  indicative  of  this  regurgitation.  Under 
these  conditions,  the  pancreatic  juice  is  present  in  amounts  sufficient  to 
have  a  considerable  proteolytic  and  fat-splitting  action. 

Time  Occupied  in  Gastric  Digestion. — Under  ordinary  conditions, 
from  three  to  four  hours  may  be  taken  as  the  average  time  occupied  by  the 
digestion  of  a  meal  in  the  stomach.  But  many  circumstances  will  modify 
the  rate  of  gastric  digestion.  The  chief  are:  The  nature  of  the  food  taken 
and  its  quantity  (the  stomach  should  be  fairly  filled,  not  distended) ;  the  time 
that  has  elapsed  since  the  last  meal,  which  should  be  at  least  enough  for  the 
stomach  to  be  quite  clear  of  food;  the  amount  of  exercise  previous  and 
subsequent  to  a  meal  (gentle  exercise  being  favorable,  overexertion  injurious, 
to  digestion);  the  state  of 'mind;  and  the  bodily  health. 

Summary  of  Changes  in  the  Food  in  Gastric  Digestion. — Briefly 
summarizing  the  action  of  gastric  juice,  the  facts  appear  as  follows:  i 
Gastric  juice  has  a  specific  digestive  action  on  protein  foods  of  all  kinds, 
converting  them  into  the  more  soluble  proteases  and  peptones.  The  action 
is  due  to  an  enzyme,  pepsin,  acting  in  and  with  an  acid,  hydrochloric  acid. 
2.  The  lipase  in  gastric  juice  produces  a  small  amount  of  fat  cleavage,  tend- 
ing to  convert  the  fats  into  fatty  acids  and  glycerin  in  which  condition  they 
are  absorbed.  The  presence  of  fat  tends  to  inhibit  the  gastric  digestion  of 
proteins.  3.  Milk  is  first  coagulated  by  a  special  enzyme,  rennin,  and  then 
digested  as  any  other  protein.  4.  Gastric  juice  dissolves  soluble  substances 
like  salts,  saccharides,  etc. 

5.  The  enzyme,  ptyalin,  continues  the  digestion  of  the  carbohydrates  in 
the  stomach  so  long  as  the  food  remains  neutral  or  alkaline,  but  they  are 
not  digested  under  the  influence  of  any  gastric  enzymes.  However,  maltase 
is  present  in  the  gastric  juice  and  aids  in  the  last  step  in  carbohydrate  hydroly- 
sis. It  is  significant  that  outside  the  body  digestion  takes  place  best  at  the 
temperature  of  the  body,  is  destroyed  by  high  heat  and  suspended  by  cold, 
o°  C.  Putrefaction  is  prevented  by  the  acid  of  natural  gastric  juice. 

MOVEMENTS  OF  THE  STOMACH. 

Attention  has  been  called  to  the  fact  that  the  stomach  is  a  muscular  sac 
capable  of  holding  quite  a  large  mass  of  food.  During  a  full  meal  as  much 
as  one  to  two  liters  or  more  of  semi-solid  food  is  packed  away  in  the  organ  in 
a  comparatively  short  space  of  time.  The  gastric  juice  is  secreted  by  the 
mucous  membrane  which  surrounds  the  surface  of  the  food  mass.  The  result 
is  that  the  secretion  begins  to  soften  and  digest  the  food  over  its  surface,  thus 
tending  to  liquefy  and  erode  away  layer  after  layer  of  the  food  mass.  The 
picture  is  made  clearer  if  one  remembers  that  the  food  mass  is  retained  al- 
most wholly  in  the  fundus  of  the  stomach.  The  pyloric  portion  of  the  stom- 


374  FOOD   AND   DIGESTION 

ach  is  quite  strongly  muscular  and  quite  definitely  marked  off  by  the  strong 
transverse  band  at  its  union  with  the  fundus. 

Acid  Closure  of  the  Cardiac  and  Pyloric  Orifices. — The  gastric 
juice  is  assisted  in  accomplishing  digestion  by  the  movements  of  the  stomach 
itself.  When  digestion  is  not  going  on,  the  stomach  is  uniformly  contracted, 
its  orifices  not  more  firmly  than  the  rest  of  its  walls;  but,  if  examined  shortly 
after  the  introduction  of  food,  it  is  found  closely  encircling  its  contents,  and 
its  orifices  are  firmly  closed  like  sphincters.  The  cardiac  orifice,  every  time 
food  is  swallowed,  opens  to  admit  its  passage  to  the  stomach,  and  immedi- 
ately closes  again.  This  closure  of  the  cardiac  orifice  is  accomplished  by  a 
local  reflex.  The  stimulus  is  the  acid  secretion  covering  the  mucous  mem- 
brane in  the  immediate  neighborhood. 

At  the  taking  of  food  or  immediately  thereafter  the  content  of  the  stomach 
begins  to  pass  through  the  pyloric  orifice  into  the  intestine.  But  the  pylorus 
is  quickly  closed  so  completely  that  Little  of  the  contents  escape  at  this 
time.  The  pylorus  is  automatically  regulated  as  demonstrated  by  Cannon. 
The  acid  gastric  content  in  the  duodenum  sets  up  a  local  reflex  that  closes 
the  pylorus  until  the  bile  and  pancreatic  juice  have  neutralized  the  acid. 
When  an  alkaline  reaction  is  produced  the  pylorus  relaxes  and  at  the  next 
peristaltic  wave  of  contraction  is  opened  again.  Indeed  it  is  claimed  that  in 
the  human  the  pylorus  takes  more  or  less  part  in  each  peristaltic  wave  passing 
from  the  stomach  on  over  the  duodenum. 

The  Peristalsis  of  the  Stomach. — The  char- 
acter of  stomach  movements  has  been  admirably 
determined  by  recent  observers  using  the  X-ray 
method.  Thus  Cannon,  working  with  cats,  has 
shown  that  in  from  five  to  ten  minutes  after  a  meal 
slight  rings  or  constrictions  occur  in  the  pyloric 
antrum  and  travel  slowly  toward  the  pyloric  sphinc- 
ter in  the  form  of  a  peristaltic  wave.  Successive 
waves  begin  a  little  further  back  toward  the  fundus 
each  time  and  follow  over  the  pyloric  antrum  with 

clock-like  regularity,  in  the  cat  one  wave  in  ten 
FIG.      261. — Diagram     to  .  . 

show  the  movement  of  food   seconds,  which  requires  in  each  case  about  twenty 

in  the  pylorus  at  times  when   seconds  for  its  completion.       In  man  they   are 
doubtless  slower.    These  peristalses  continue  dur- 
ing the  whole  period  of  digestion  for  as  much  as  seven  or  even  more  hours. 

These  peristaltic  contractions  aid  the  gastric  juice  in  carrying  away  the 
softened  layers  of  food  by  propelling  it  into  the  pylorus.  There  it  is  thoroughly 
mixed  with  the  gastric  juice,  forming  the  chyme.  Figure  261  gives  an  idea  of 
the  movement  of  the  food  in  the  antrum.  The  peristaltic  contractions  carry 
it  forward,  but  if  the  valve  does  not  open  to  permit  passage  to  the  duodenum, 
then  the  pressure  will  force  the  chyme  back  through  the  center  toward  the 


THE   PERISTALSIS    OF   THE    STOMACH  375 

fundus.  After  several  minutes,  i.e.,  when  the  secretion  of  alkaline  bile  and 
pancreatic  juice  into  the  duodenum  is  well  established,  the  pyloric  sphincter 
will  relax  more  often  to  allow  fluid  food  to  pass  to  the  duodenum,  but  when 
the  more  solid  particles  come  up  against  the  sphincter  it  promptly 
contracts  and  remains  so  for  some  time.  Toward  the  completion  of  digestion 
even  solid  undigested  particles  are  carried  on  into  the  intestine. 

Hunger  Contraction.  —  Our 
present  knowledge  of  the  charac- 
ter and  nervous  regulation  of  the 

peristaltic      contractions      of      the      f     ]^^  J     ''A.M. 

stomach  has  been  recently  immeas- 
urably advanced  by  the  work  of 
Carlson.  His  studies  have  been  on 
man,  with  and  without  gastric 
fistula,  and  on  dogs.  As  a  result 
there  can  no  longer  be  doubt  that 

the  sensation  of  hunger  is  attended        (      ^        /  ]      12  M . 

with  characteristic  contractions  of 
the  empty  stomach.  Carlson's 
studies  of  the  contractions  of  the 
empty  stomach  has  established  a 
number  of  points  in  the  nervous 

control.      For  example,  the  vagus 

,  .    .  a 

nerves  have  a  tonic  influence  over 

the  organ,  and  their  section  "  leaves 

the  empty  stomach  on  the  whole 

permanently    hypotonic."        This 

control    does   not  readily  yield  to 

the  usual  reflexes.     The  splanchnic 

nerves  are,  on  the  other  hand,  in-        (       )     ^/  5P.M. 

hibitory  for  the   stomach.      It    is 

through    this    channel    that     the 

psychic    and    other  reflexes  act  to       FlG>   262.— Outlines  of  the    Roentgen-ray 

control  the  organ.    Carlson  studied    Shadows  of  the  Stomach  Content  as  Digestion 

.    ,      ,    Progresses.     (Cannon.) 
the  phenomena  in  dogs  which  had 

both  the  vagi  and  the  splanchnics  sectioned.  The  stomach  thus  isolated 
manifests  the  usual  hunger  peristalses  and  often  "the  hunger  contractions 
are  identical  in  rate  and  character  with  those  of  the  intact  stomach  in  normal 
(strong)  tonus."  This  indicates  that  "the  primary  stimulus  to  these  con- 
tractions is  not  to  be  sought  in  the  extrinsic  nerves."  It  follows  therefore 
that  the  extrinsic  nerves  are  merely  regulative  and  modifying  for  the  con- 
tractions of  the  otherwise  automatic  organ.  Of  the  two  mechanisms,  the 
vagus  is  the  more  vital  and  least  readily  disturbed  in  its  control. 


376 


FOOD   AND   DIGESTION 


FIG.  263. — Horizontal  Section  of  a 
Small  Fragment  of  the  Mucous  Mem- 
brane, including  one  entire  crypt  of 
Lieberkuhn  and  parts  of  several  others. 


Vomiting.  —  The  expulsion  of  the  contents  of  the  stomach  in  vomiting 
is  preceded  by  a  deep  inspiration  with  closure  of  the  glottis,  followed  im- 
mediately afterward  by  strong  contractions  of  the  muscles  of  the  abdomen, 
diaphragm,  and  stomach.  The  diaphragm  forms  an  unyielding  surface 
against  which  the  stomach  can  be  pressed.  In  this  way  as  well  as  by 

its  own  contraction  the  diaphragm  is 
fixed,  to  use  a  technical  phrase.  At  the 
same  time  the  cardiac  sphincter  muscle  is 
relaxed,  and  the  orifice  which  it  naturally 
guards  is  actively  dilated.  The  pylorus 
is  closed  and,  the  stomach  itself  also  con- 
tracting, the  action  of  the  abdominal 
muscles  produces  strong  compression 
which  expels  the  contents  of  the  organ 
through  the  esophagus,  pharynx,  and 
mouth.  Reversed  peristaltic  action  of  the 
esophagus  probably  increases  the  effect. 
It  has  been  frequently  stated  that  the 
stomach  itself  is  quite  passive  during 
vomiting,  and  that  the  expulsion  of  its 

contents  is  effected  solely  by  the  pressure  exerted  upon  it  when  the  capacity 
of  the  abdomen  is  diminished  by  the  contraction  of  the  diaphragm.  It  is 
true  that  facts  are  wanting  to  demonstrate  with  certainty  the  contraction 
of  the  stomach  in  vomiting;  but  cases  of  fistulous  opening  into  the  organ 
appear  to  support  the  belief  that  it  does  take  place;  and  the  analogy  of  the 
case  of  the  stomach  with  that  of  the  other  hollow  viscera,  as  the  rectum 
and  bladder,  may  also  be  cited  in  confirmation. 

Vomiting  is  a  reflex  act.  It  can  be  excited  by  irritation  of  the  lining  of 
the  stomach,  which  is  perhaps  the  normal  stimulus.  It  is  excited  by  stimula- 
tion or  irritation  of  other  parts  of  the  alimentary  tube;  i.e.,  the  pharynx,  the 
uvula,  the  intestine,  etc.  Vomiting  may  occur  from  stimulation  of  sensory 
nerves  from  many  organs,  e.g.,  kidney,  testicle,  etc.,  or  by  impulses  arising 
in  the  organs  of  special  sense,  the  eye,  olfactory  membrane,  etc.  The  sensory 
impulses  are  co-ordinated  by  a  nerve  center  located  in  the  medulla.  The 
center  may  also  be  stimulated  by  impressions  from  the  cerebrum  and  cere- 
bellum or  by  changes  arising  in  the  center  itself,  the  so-called  central  vomiting 
occurring  in  disease  of  those  parts.  The  efferent  impulses  are  carried  by  the 
phrenics  and  other  spinal  nerves  and  by  the  vagus. 

DIGESTION  IN  THE  INTESTINES. 

The  food  that  enters  the  small  intestine  has  already  been  subjected  to  two 
digestive  enzymes.  The  ptyalin  of  the  saliva  and  the  pepsin  of  the  gastric 
juice  together  with  the  mechanical  processes  involved  have  reduced  the  food 


THE  PANCREAS 


377 


to  a  pulpy  mass,  the  chyme.     This  peptonized/00d  contains  most  of  the  total 
quantity  of  food  eaten,  little  having  been  absorbed,  as  we  shall  see  later,  but 


FIG.  264.  FIG.  265. 

FIG.  264. — Piece  of  Small  Intestine  (previously  distended  and  hardened  by  alcohol), 
Laid  open  to  Show  the  Normal  Position  of  the  Valvulae  Conniventes. 

FIG.  265. — Section  of  the  Pancreas  of  a  Dog  During  Digestion,  a,  Alveoli  lined  with 
cells,  the  outer  zone  of  which  is  well  stained  with  hematoxylin;  d,  intermediary  duct  lined 
with  squamous  epithelium.  X  350.  (Klein  and  Noble  Smith.) 

much  of  the  starch  has  been  changed  to  soluble  maltose  and  dextrose  and 
the  protein  to  albumoses  and  peptones.  The  discharge  from  the  stomach 
through  the  pyloric  valve  to  the  duo- 
denum has  been  going  on  through 
three  or  four  hours  on  an  average  for 
each  full  meal.  This  stream  of  food 
passing  down  the  small  intestine, 
slowly  because  of  the  valvulse  con- 
niventes,  meets  a  number  of  secretions 
which  contain  enzymes  which  act  on 
each  of  the  three  great  food  principles, 
proteins,  fats,  and  carbohydrates. 
These  secretions  are  the  pancreatic 
fluid,  the  succus  entericus,  and  the 
bile. 

The  Pancreas. — The  pancreas 
is  situated  within  the  curve  formed 
by  the  duodenum,  and  its  main  duct 
opens  into  that  part  of  the  small  intes- 
tine through  a  duct  common  to  it  and 

to  the  liver  and  about  two  and  a  half  , 

FIG.    266. — Section  of   the  Pancreas  of 

inches  from  the  pylorus.  Armadillo,   Showing   the  Two   Kinds  of 

The  pancreas  bears  some  resem-  Gland-structure.    (V.  D.  Harris.) 


378 


FOOD   AND    DIGESTION 


blance  in  structure  to  the  salivary  glands.  Its  capsule  and  septa,  as  well  as 
the  blood  vessels  and  lymphatics,  are  similarly  distributed.  It  is,  however, 
looser,  the  lobes  and  lobules  being  less  compactly  arranged. 

Heidenhain  has  observed  that  the  alveolar  cells  in  the  pancreas  of  a  fast- 
ing dog  consist  of  two  zones,  an  inner  or  central  zone  which  is  finely  granular, 
.and  which  stains  feebly,  and  a  smaller  parietal  zone  of  finely  striated  proto- 


FIG.  267.— Duct  with  Laterals  to  the  Alveoli.     Silver  method  of  Golgi  (E.  Muller).     A 
Duct  with  branches;  m,  between  the  cells.     B,  Laterals  more  strongly  magnified. 

plasm  which  stains  easily.  The  nucleus  is  partly  in  one,  partly  in  the  other 
zone.  During  secretion  it  is  found  that  the  outer  zone  increases  in  size,  and 
the  central  granular  zone  diminishes,  as  in  the  case  of  the  salivary  glands. 
The  pancreatic  cell  itself  becomes  smaller  from  the  discharge  of  the  secretion. 
During  a  period  of  rest  the  granular  zone  again  increases  in  size  and  the 
outlines  of  the  cells  become  full  and  indistinct.  The  granules,  as  in  the  sali- 
vary cells,  are  the  material  from  which,  under  certain  conditions,  the  fer- 
ments of  the  gland  are  developed,  and  which  are  therefore  a  zymogen.  In 
addition  to  the  ordinary  alveoli  of  the  pancreas  there  are  distributed  irregu- 


THE    PANCREATIC   JUICE 


379 


larly  in  the  gland  other  collections  of  cells  of  a  different  character,  the  islands 
of  Langerhans.  These  cells  are  considerably  smaller,  their  protoplasm  is 
more  granular  and  less  easily  stained  with  hematoxylin,  and  their  nuclei  are 
small  and  stain  deeply.  The  collections  of  cells  vary  in  size  and  shape. 
The  islands  of  Langerhans'  cells  are  not  concerned  with  the  production  of 
the  pancreatic  juice.  The  special  form  of  nerve  terminations,  called 
Pacinian  corpuscles,  are  often  found  in  the  pancreas. 

The  secretion  of  the  pancreas  has  been  obtained  for  purposes  of  experi- 
ment from  the  lower  animals  and  from  man  in  at  least  one  case.  A 
pancreatic  fistula  is  established  in  the  dog  by  opening  the  abdomen  and 
exposing  the  duct  of  the  gland  which  is  then  made  to  communicate  with 
the  exterior.  In  Pawlow's  method  a  circular  bit  of  the  intestinal  mucous 
membrane  around  the  mouth  of  the  duct  in  the  intestine  is  brought  to 
the  surface  and  stitched  into  the  wound.  The  secretion  is  then  easily  col- 
lected into  a  vessel  suspended  under  the  opening. 

The  Pancreatic  Juice. — Pancreatic  juice  is  colorless,  transparent, 
slightly  viscid,  and  alkaline  in  reaction.  It  varies  in  specific  gravity  from 
i .  oio  to  i .  030,  according  as  it  is  obtained  from  a  permanent  fistula,  when  it  is 
more  watery,  or  from  a  newly  opened  duct.  The  solids  vary  in  a  temporary 
fistula  from  80  to  100  parts  per  thousand,  and  in  a  permanent  one  from  16  to 
50  per  thousand.  It  is  characterized  by  having  three  distinct  and  important 
enzymes  known  as  trypsin,  amylopsin,  and  steapsin,  whose  actions  are,  respect- 
ively, proteolytic,  amylolytic,  and  lipolytic  (fat-splitting).  Maltase,  which 
inverts  the  disaccharides,  is  also  present,  and  a  rennin  is  found  in  the  pan- 
creatic juice. 

CHEMICAL  COMPOSITION  OF  PANCREATIC  JUICE.      (C.  SCHMIDT.) 


From  a  dog. 

Recent 
fistula. 

Perma- 
nent 
fistula. 

Water 

ooo  .  76 

080  .4  <; 

'Solids 

OQ  .  24 

IQ  .  c  c 

Organic  substances                   .             .           

QO  .44 

12.71 

Ash 

8.80 

6.84 

Sodium  carbonate 

o.  t:8 

7       71 

Sodium  chloride                        .             

7  .  s  e 

2  .  <O 

Calcium,  magnesium,  and  sodium  phosphates    .... 

o-53 

0.08 

An  extract  of  pancreas  made  from  the  gland  which  has  been  removed 
from  an  animal  killed  during  digestion  possesses  the  active  properties  of 
pancreatic  secretion.  It  is  made  by  first  dehydrating  in  absolute  alcohol 
the  gland  which  has  been  cut  up  into  small  pieces.  After  the  entire  removal 


FOOD   AND   DIGESTION 


of  the  alcohol  the  gland  is  pulverized  and  extracted  in  strong  glycerin. 
The  amount  of  the  ferment  greatly  increases  if  the  gland  be  exposed  to  the 
air  for  three  or  four  hours  before  placing  in  alcohol;  indeed,  a  glycerin 
extract  made  from  the  gland  immediately  upon  the  removal  from  the  body 
often  appears  to  contain  none  of  the  ferments.  The  conversion  of  zymogen 
in  the  gland  into  the  ferment  takes  place  only  after  the  gland  stands  a  while. 


1. 


z. 


3. 


s. 


6. 


7. 


. 


Jit 


\ 


32 


U 


16 


JtCOk. 


-  (Stead 


FIG.  268. — Three  Curves  Showing  the  Secretion  of  Pancreatic  Juice  upon  a  Diet  (i) 
of  600  cc.  of  milk;  (2)  of  250  gm.  of  bread;  (3)  of  100  gm.  of  meat.  The  divisions  along 
the  abscissae  represent  hours  after  the  beginning  of  the  meal;  the  figures  along  the  ordinates 
represent  the  quantity  of  the  secretion  in  cubic  centimeters.  (Walter.) 

Dilute  acid  assists  or  accelerates  the  conversion,  and  if  a  recent  pancreas  be 
rubbed  up  with  dilute  acid  before  dehydration,  a  glycerin  extract  made 
afterward,  even  though  the  gland  may  have  been  only  recently  removed  from 
the  body,  is  very  active. 

Nervous  Regulation  of  the  Secretion  of  the  Pancreas. — Fibers  from 
the  vagus  and  from  the  splanchnics  are  distributed  to  the  pancreas.  In 
Pawlow's  laboratory  it  has  been  found  that  stimulation  of  these  nerves  leads 
to  the  increased  secretion  of  the  pancreas.  Popielski,  in  studying  the  effects 


ACTION  OF  THE  ENZYMES  OF  PANCREATIC  JUICE       381 

of  dilute  hydrochloric  acid  solution  in  the  duodenum,  which  resulted  in  a 
marked  increase  of  pancreatic  secretion,  explained  the  phenomenon  as  a 
local  nerve  reflex. 

Doubt  has  been  cast  on  the  whole  question  of  nervous  control  by  the 
recent  discovery  of  the  fact  that  acid  (o .  4  per  cent,  hydrochloric  acid)  in  the 
duodenum  results  in  the  production  of  a  chemical  substance,  by  the  duodenal 
mucous  membrane.  This  substance  secretin,  is  absorbed  into  the  circulation 
and  acts  specifically  on  the  pancreas  to  produce  increased  activity  of  the  pan- 
creatic cells.  Acid  extracts  of  the  duodenal  mucous  membrane  produce  the 
same  effects  on  the  pancreas,  in  fact  this  is  the  current  method  of  experiment- 
ally stimulating  the  flow  of  pancreatic  juice  at  the  present  time,  the  secretion 
being  collected  from  a  tube  introduced  into  the  duct. 

Under  the  normal  stimulus  of  food,  the  flow  of  pancreatic  juice  is  greatly 
increased.  The  increase  continues  to  a  maximum  in  from  two  to  three  hours, 
after  which  it  gradually  decreases  through  the  period  of  digestion.  Pawlow 
has  found  a  certain  amount  of  adaptation,  not  only  of  the  quantity  but  of 
the  enzyme  composition  of  the  pancreatic  secretion,  to  the  kind  and  character 
of  the  food  (in  dogs). 

Action  of  the  Enzymes  of  Pancreatic  Juice. — The  secretion  of  the 
pancreas  accomplishes  its  digestive  action  by  means  of  the  enzymes  given 
above,  viz.,  trypsin,  amylopsin,  steapsin,  and  maltase. 

Trypsin. — Trypsin  is  a  proteolytic  enzyme.  Strange  to  say,  it  does  not 
exist  in  the  fresh  pancreatic  juice  as  such,  but  makes  its  appearance  only 
when  there  is  an  admixture  with  the  succus  entericus,  the  secretion  of  the 
mucous  membrane  of  the  intestine.  The  succus  entericus  contains  an  ac- 
tivating enzyme,  enterokinase,  which  converts  the  inactive  and  stable  trypsin- 
ogen  of  the  pancreatic  juice  into  the  active  but  less  stable  trypsin.  This  fact 
is  another  of  the  wonderful  series  of  contributions  to  the  exact  knowledge 
of  the  subject  of  digestion  made  from  Pawlow's  laboratories. 

Trypsin,  like  pepsin-hydrochloric  acid,  converts  proteins  into  proteoses 
and  peptones.  The  change,  however,  does  not  stop  here;  the  hydrolysis 
with  trypsin  goes  much  further.  While  simple  amino-acids,  with  the  excep- 
tion of  traces  of  tyrosine,  are  not  found  in  gastric  digestions,  these  are  rapidly 
split  off  in  the  tryptic  cleavage.  Thus  in  tryptic  digestion  are  formed:  tyro- 
sine,  leucine,  cystine,  amino-valerianic  acid,  asparaginic  acid,  glutaminic 
acid,  histidine,  lysine,  and  arginine.  A  portion  of  the  protein,  however, 
is  not  completely  broken  down,  the  residue  consisting  of  polypeptids 
containing  proline  and  phenyl-alanine  combined  with  small  amounts  of 
alanine,  leucine,  aspartic  acid,  and  glutaminic  acid.  Glycocoll,  when  pres- 
ent in  the  protein  digested,  is  also  combined  in  the  resistant  polypeptids. 
Crystals  of  leucine  and  of  tyrosine,  especially,  can  be  found  in  tryptic  diges- 
tion mixtures.  The  products  formed  from  protein  in  tryptic  digestion  may 
be  given  in  the  following  graphic  scheme: 


382  FOOD   AND    DIGESTION 

Protein 
Proteoses 
Peptones 


Polypeptids  Amino-acids 

Combinations  of  proline,  phenyl-  Tyrosine,     tryptophane,     cystine, 

alanine  and  glycocoll,  with  rela-  alanine,    amino-valerianic   acid, 

tively  small  amounts  of  alanine,  leucine,    aspartic    acid,  glutam- 

leucine,    asparaginic   acid,    and  inic  acid,  histidine,  lysine,  argi- 

glutaminic  acid.  nine. 

The  ferment  trypsin  acts  best  in  an  akaline  medium,  but  will  act  also 
in  a  neutral  medium,  or  in  the  presence  of  a  very  small  amount  of  combined 
acid;  it  will  not  work  in  the  presence  of  free  acid.  It  therefore  differs  from 
pepsin  in  being  able  to  act  without  the  aid  of  any  other  substance  than  water. 
In  the  process  of  tryptic  digestion,  protein  matter  does  not  swell  up  at  first,, 
but  seems  to  be  corroded  at  once. 

Amylopsin. — Starch  is  converted  by  amylopsin  into  maltose  by  hydro- 
lytic  action  similar  to  that  of  ptyalin,  ery thro- dextrin  and  one  or  more 
achroo-dextrins  being  the  intermediate  products.  The  amylolytic  enzyme 
of  the  pancreatic  juice,  which  cannot  be  distinguished  from  ptyalin,  is  called 
amylopsin.  The  maltose  thus  formed  is  converted  to  dextrose  by  the  maltase, 
in  which  form  it  is  ultimately  absorbed. 

Pancreatic  juice,  according  to  certain  observers,  possesses  the  property 
of  curdling  milk.  It  contains  a  special  ferment,  rennin,  for  that  purpose. 
The  ferment  is  distinct  from  trypsin,  and  will  act  in  the  presence  of  an  acid 
(W.  Roberts).  The  milk-curdling  ferment  of  the  pancreas  is,  in  some  pan- 
creatic extracts,  said  to  be  quite  powerful,  insomuch  that  i  cc.  of  a  brine  ex- 
tract will  coagulate  50  cc.  of  milk  in  a  minute  or  two. 

Steapsin  orLipase. — Oils  and  fats  are  emulsified  and  saponified  by  the  pan- 
creatic secretion.  The  terms  emulsification  and  saponification  may  need  a 
little  explanation.  The  term  emulsification  is  used  to  signify  an  important 
mechanical  change  in  oils  or  fats,  whereby  they  are  made  into  an  emulsion 
or,  in  other  words,  are  minutely  subdivided  into  small  microscopic  particles. 
If  a  drop  of  an  emulsion  be  looked  at  under  the  microscope,  an  immense 
number  of  minute  rounded  particles  of  oil  or  fat  of  varying  sizes  will  be 
seen.  The  more  complete  the  emulsion  the  smaller  are  these  particles.  In 
milk,  which  is  a  splendid  example  of  an  emulsion,  the  fat  droplets  vary  in 
diameter  between  i  and  5^.  An  emulsion  is  formed  at  once  if  oil,  which 
when  old  is  slightly  acid  from  the  presence  of  free  fatty  acid,  is  mixed 
with  an  alkaline  solution. 

Saponification  signifies  a  distinct  chemical  change  in  the  composition 
of  oils  and  fats.  An  oil  or  fat  being  made  up  chemically  of  glycerin,  a 


CONDITIONS    OF    THE    PANCREATIC   ENZYMES  383 

triatomic  alcohol,  and  three  fatty-acid  radicles  which  may  or  may  not  be 
identical,  when  an  alkali  (potassium  hydrate)  is  added  to  it  two  changes 
take  place;  first,  the  oil  or  fat  is  split  up  into  glycerin  and  its  fatty  acid;  second, 
the  fatty  acids  combine  with  the  alkali  to  form  soaps  which  are  chemically 
known  as  stearate,  oleate,  or  palmitate  of  potassium  according  to  the  particular 
fatty  acid  or  acids  involved.  Saponification  thus  means  a  chemical  splitting 
up  of  oils  or  fats  into  new  compounds,  and  emulsification  means  merely  a 
mechanical  splitting  up  into  minute  particles.  The  pancreatic  juice  has 
been  for  many  years  credited  with  the  possession  of  a  special  ferment, 
which  was  called  by  Claude  Bernard  steapsin,  and  which  is  a  lipase  or 
fat-splitting  ferment.  This  ferment  has  not  been  isolated,  but  its  pres- 
ence may  be  demonstrated  by  adding  portions  of  the  fresh  pancreas 
to  butter  or  other  fat  and  maintaining  the  proper  temperature.  Its 
action  is  made  manifest  by  the  liberation  of  butyric  acid,  which  imparts  the 
typical  odor  to  rancid  butter. 

The  older  theory  was  that  only  a  small  portion  of  the  fat  eaten  was  thus 
changed  into  soap,  and  that  the  function  of  the  saponified  fat  was  to  assist 
in  the  emulsification  of  the  remaining  major  part,  a  process  favorably 
influenced  by  the  bile.  Although  the  proper  emulsification  of  fat  is  indeed 
a  preliminary  step  favoring  more  effective  contact  of  the  fat  splitting  enzyme, 
lipase,  we  now  know  that  all  the  fat  is  dissociated  into  fatty  acid  and  glycerin 
before  abosrption  can  occur.  When  in  disease  the  entrance  of  the  pancreatic 
juice  and  the  bile  into  the  intestine  is  interfered  with,  the  feces  contain  an 
excess  of  fat. 

All  recent  experiments,  however,  tend  to  support  the  view  of  Pflliger 
that  the  entire  fat  of  the  food  is  changed  into  fatty  acids  and  glycerin;  that 
the  fatty  acids  are  entirely,  or  in  part,  changed  to  soaps;  and  that  these  soaps, 
or  mixture  of  soaps  and  free  fatty  acids,  are  absorbed  in  solution.  The 
chief  facts  favoring  this  view  are  that:  (i)  The  reaction  of  lipase  is  sufficiently 
rapid  to  allow  the  saponification  of  a  full  fatty  meal  within  the  ordinary 
period  of  digestion;  (2)  histological  examination  has  never  shown  that  fat 
particles  can  pass  into  a  columnar  cell,  and  droplets  have  not  been  found  in  the 
broad  striated  border  of  the  cell;  (3)  the  fat  globules  found  in  columnar  cells 
after  a  fatty  meal  grow  steadily  larger  as  the  period  of  absorption  progresses, 
indicating  that  they  are  deposited  from  solution;  (4)  the  fatty  acids  are  easily 
soluble  in  bile  solutions,  and  the  solubility  of  the  soaps  is  greatly  increased 
by  the  presence  of  bile.  The  fat  constituents,  according  to  this  theory,  are 
recombined  in  the  columnar  cells  to  form  neutral  fats  where  their  presence 
can  be  easily  demonstrated  by  methods  of  staining. 

Conditions  which  Influence  the  Action  of  the  Pancreatic  Enzymes. 
— The  various  pancreatic  enzymes  are  influenced  by  heat,  by  the  presence  of 
an  excess  of  digestion  products,  etc.,  in  the  same  way  as  ptyalin  and  pepsin. 
Pancreatic  enzymes  act  in  a  neutral,  but  best  in  an  alkaline  solution.  The 


384 


FOOD   AND   DIGESTION 


trypsin,  strange  to  say,  is  quickly  destroyed  by  the  alkaline  solution  (Bayliss 
and  Starling).     The  pancreatic  juice  offers  the  special  case  of  a  secretion  of 


FIG.  269.  FIG.  270. 

FIG.  269. — The  Liver  from  Below  and  Behind.  L.  S.,  Spigelian  lobe;  L.  C.,  caudate 
lobe;!,.  Q.,  quadrate  lobe;  R.L.,  right  lobe;Z.L.,  left  lobe;  g.  bl.,  gall-bladder;  v.c.i.,  inferior 
vena  cava;  u.f.,  umbilical  fissure;  /.  d.  v.,  fissure  of  the  ductus  venosus;  p,  portal  fissure  with 
portal  vein,  hepatic  artery,  and  bile-duct.  (Wesley,  from  a  His  model.) 

FiG.  270. — Portion  of  a  Lobule  of  Liver,  a,  Bile  capillaries  between  liver  cells,  the 
network  in  which  is  well  seen;  b,  blood  capillaries.  X  350.  (Klein  and  Noble  Smith.) 

proenzyme  which  is  stable  in  alkaline  solution  until  acted  on  by  enterokinase. 
The  amount  of  kinase  present  will,  therefore,  markedly  influence  the 
amount  of  digestion  of  protein  per  unit  of  time. 


FIG.  271. — Hepatic  Cells  and  Bile  Capillaries,  from  the  Liver  of  a  Child  Three  Months 
Old.  Both  figures  represent  fragments  of  a  section  carried  through  the  periphery  of  a 
lobule.  The  red  corpuscles  of  the  blood  are  recognized  by  their  circular  contour;  vp 
corresponds  to  an  interlobular  vein  in  immediate  proximity  with  which  are  the  epithelial 
cells  of  the  biliary  ducts.  (E.  Hering.) 

The  Secretions  of  the  Liver. — The  liver,  the  largest  gland  in  the  body, 
situated  in  the  abdomen  on  the  right  side  chiefly,  is  an  extremely  vascular 
organ,  and  receives  its  supply  of  blood  from  two  distant  sources,  viz.,  from 
the  portal  vein  and  from  the  hepatic  artery,  while  the  blood  is  returned  from 
it  into  the  vena  cava  inferior  by  the  hepatic  veins.  Its  secretion,  the  bile, 


STRUCTURE    OF    THE    LIVER 


385 


is  conveyed  from  it  by  the  hepatic  duct,  either  directly  into  the  intestine  or 
when  digestion  is  not  going  on,  into  the  cystic  duct,  and  thence  into  the  gall- 
bladder, where  it  accumulates  until  required.  The  portal  vein,  hepatic 
artery,  and  hepatic  duct  branch  together  throughout  the  liver,  while  the 
hepatic  veins  and  their  tributaries  run  by  themselves.  The  interstices  of  the 
vessels  are  filled  by  the  liver  cells. 

Structure  of  the  Liver. — The  liver  is  made  up  of  small  roundish  or 
oval  portions  called  lobules,  each  of  which  is  about  ^  of  an  inch  (about 


P  x 


FIG.   272.—  Section  of  Liver.     X  80.     P,  Portal  vein;  H,  hepatic  artery;  B,  bile-duct. 

(Hendrickson.) 

i  mm.)  in  diameter,  and  includes  the  minute  hepatic  artery  and  hepatic 
duct.  The  hepatic  cells,  which  form  the  glandular  or  secreting  part  of  the 
liver,  are  of  spheroidal  form,  somewhat  polygonal  from  mutual  pressure, 
about  25  to  3o//  in  diameter,  and  possess  one,  sometimes  two  nuclei.  The 
cell  substance  contains  a  variable  amount  of  glycogen  and  often  some  fatty 
globules  and  possibly  some  granules  of  bile  pigment. 

The  bile  capillaries  commence  between  the  hepatic  cells,  and  are  bounded 
by  a  delicate  membranous  wall  of  their  own.  They  appear  to  be  always 
bounded  by  hepatic  cells  on  all  sides,  and  are  thus  separated  from  the  nearest 
blood  capillary  by  at  least  the  breadth  of  one  cell,  figures  271  and  272. 


386  FOOD   AND   DIGESTION 

The  gall-bladder,  g.  bl,  figure  269,  is  a  pyriform  sac  attached  to  the  under 
surface  of  the  liver,  and  supported  also  by  the  peritoneum.  The  larger  end, 
or  fundus,  projects  beyond  the  front  margin  of  the  liver,  while  the  smaller 
end  contracts  into  the  cystic  duct.  It  is  a  muscular  walled  reservoir  covered 
with  a  serous  epithelium  and  lined  by  mucous  membrane.  The  function 
of  the  gall-bladder  is  to  retain  the  bile  during  the  interval  of  digestion. 

The  Bile. — The  bile  is  a  somewhat  viscid  fluid,  of  a  yellow,  reddish- 
yellow,  or  green  color,  a  strongly  bitter  taste,  and,  when  fresh,  with  a  scarcely 
perceptible  odor;  it  has  a  neutral  or  slightly  alkaline  reaction,  and  its  specific 
gravity  is  about  1.020.  Its  color  and  consistency  vary  much,  quite  inde- 
pendent of  disease;  but,  as  a  rule,  bile  becomes  gradually  more  deeply  colored 
and  thicker  as  it  advances  along  its  ducts,  or  when  it  remains  long  in  the  gall- 
bladder where  it  becomes  more  viscid  and  ropy,  darker,  and  more  bitter. 
This  is  on  account  of  its  greater  degree  of  concentration,  from  resorption  of 
its  water,  and  also  from  being  mixed  with  mucus,  lipoids,  and  phosphatid 
proteins  secreted  by  the  lining  membrane  of  the  gall-bladder. 

CHEMICAL  COMPOSITION  OF  HUMAN   BILE.     (FRERICHS.) 

Water 859.2 

Solids  — Bile  salts 91.5 

Fat 9.2 

Cholesterol 2.6 

Proteins  and  coloring  matters 29.8 

Salts 7.7 

140. 8 


1000.0 

Bile  salts  can  be  obtained  as  colorless,  exceedingly  deliquescent  crystals, 
soluble  in  water,  alcohol,  and  alkaline  solutions,  giving  to  the  watery  solution 
the  taste  and  general  characters  of  bile.  They  consist  of  sodium  salts  of  gly- 
cocholic  and  taurocholic  acids;  the  formula  of  the  former  being  C26H42NaNO6, 
and  of  the  latter  C26H44NaNSO7. 

The  bile  acids  are  easily  decomposed  by  the  action  of  dilute  acids  or  alkalies 
thus: 

C26H43N06  +  H20  =  C24H4006  +  CH2.NH2.COOH. 
Glycocholic  Acid  Cholic  Acid     Glycocoll 

C26H46NSO7  +  H2O  =  C24H40O5  +  CH2.NH2.CH2SO3H. 
Taurocholic  Acid  Cholic  Acid       Taurine 

Glycocoll  is  amido-acetic  acid,  i.e.,  acetic  acid,  CH3COOH  with  one  of  the  atoms  of  H 
replaced  by  the  radical  amidogen  CH2.NH2.COOH.  Taurine  likewise  is  amino-ethyl- 
sulphonic  acid.  Accordingly,  it  has  the  formula  CH2NH2CH2SO3H.  The  proportion  of 
these  two  salts  in  the  bile  of  different  animals  varies,  e.g.,  in  the  ox  bile  the  glycocholate  is  in 
great  excess,  whereas  the  bile  of  the  dog,  cat,  bear,  and  other  carnivora  contains  taurocho- 
late  alone.  In  human  bile  the  glycocholate  is  in  excess  (4.8  to  1.5). 

The  yellow  coloring  matter  of  the  bile  of  man  and  the  carnivora  is  termed 


THE   BILE 


387 


bilirubin,  C16H18N2O3,  is  crystallizable  and  insoluble  in  water,  and  soluble  in 
chloroform  or  carbon  disulphide.  A  green  coloring  matter,  biliverdin, 
C16H18N2O4,  which  always  exists  in  large  amount  in  the  bile  of  herbivora,  is 
formed  from  bilirubin  on  exposure  to  the  air  or  by  subjecting  the  bile  to  any 
other  oxidizing  agency,  as  by  adding  nitric  acid.  Biliverdin  is  soluble  in 
alcohol,  glacial  acetic  acid,  and  strong  sulphuric  acid,  but  insoluble  in  water, 
in  chloroform,  and  ether.  It  is  usually  amorphous,  but  may  sometimes 
crystallize  in  green  rhombic  plates. 

There  is  a  close  relationship  between  the  coloring  matters  of  the  blood 
and  of  the  bile  and,  it  may  be  added,  between  these  and  that  of  the  urine, 
urobilin,  and  of  the  feces,  stercobilin.     It  is  probable  they  are,  all  of  them, 
varieties  of  the  same  pigment,  or  derived  from 
the  same    source.      Cholesterol    C27H45OH, 
and  lecithin,  C43H84NPO8  are  constant  con- 
stituents of  bile.      Iron  is  found  among  the 
salts  of  the  ash. 

The  Role  of  Bile  in  Intestinal  Digestion. 
—Though  it  is  not  a  true  digestive  fluid,  in 
that  it  has  no  ferment  and  digests  nothing 
itself,  yet  it  must  be  regarded  as  an  important 
aid  to  digestion  for  the  following  reasons:  (a) 
Bile  assists  in  emulsifying  the  fats  of  the  food, 
and  dissolves  the  fatty  acids  thus  rendering 
them  more  capable  of  absorption.  For  it  has 
appeared  in  some  experiments  in  which  the  common  bile-duct  was  tied  that, 
although  the  process  of  digestion  in  the  stomach  was  unaffected,  chyle 
was  no  longer  well  formed.  The  contents  of  the  lacteals  consisted  of  clear, 
colorless  fluid,  instead  of  being  opaque  and  white,  as  they  ordinarily  are 
after  feeding.  It  is,  however,  the  combined  action  of  the  bile  with  the 
pancreatic  juice  to  which  the  emulsification  is  due  rather  than  to  that  of  the 
bile  alone.  The  bile  itself  has  a  very  feeble  emulsifying  power.  If  the 
theory  be  accepted  that  fats  are  absorbed  as  fatty  acids  and  soaps,  in 
solution,  the  action  of  the  bile  becomes  very  important  because  solutions  of 
bile  salts  have  the  power  of  dissolving  the  fatty  acids.  The  moistening  of 
the  mucous  membrane  of  the  intestines  with  bile,  for  this  very  reason, 
facilitates  absorption  of  fatty  matters  through  it. 

(6)  The  bile,  like  the  gastric  fluid,  has  a  certain  but  not  very  considerable 
antiseptic  power,  and  may  serve  to  prevent  the  decomposition  of  food  during 
the  time  of  its  sojourn  in  the  intestines.  Experiments  show  that  the  contents 
of  the  intestines  are  much  more  fetid  after  the  common  bile-duct  has  been 
tied  than  at  other  times.  Moreover,  it  is  found  that  the  mixture  of  bile  with 
a  fermenting  fluid  stops  the  process  of  fermentation.  Contact  with  bile 
also  destroys  the  digestive  action  of  pepsin  on  protein. 


FIG.  273. — Crystalline  Scales  of 
Cholesterol. 


388  FOOD   AND   DIGESTION 

Bile  is  also  an  excretive  fluid  carrying  waste  products  thrown  off  by  the 
liver.  The  liver  during  fetal  life  is  proportionately  larger  than  it  is  after 
birth,  and  the  secretion  of  bile  is  active,  although  there  is  no  food  in  the  in- 
testinal canal  upon  which  it  can  exercise  any  digestive  property.  At  birth, 
the  intestinal  canal  contains  concentrated  bile,  mixed  with  intestinal  secretion 
and  this  constitutes  the  mecanium,  or  feces  of  the  fetus.  In  the  fetus,  there- 
fore, the  main  purpose  of  the  secretion  of  bile  must  be  directly  excretive. 
Probably  all  the  residue  of  the  bile  secreted  in  fetal  life  is  incorporated  in 
the  meconium,  and  with  it  discharged. 

Mode  of  Secretion  and  Discharge  of  Bile. — In  considering  the  flow 
of  bile  into  the  intestine,  two  factors  are  involved.  These  are  the  emptying 
of  the  gall-bladder  and  an  increased  secretion  by  the  hepatic  cells. 

The  secretion  oj  bile  can  be  studied  by  tying  the  common  bile-duct  of 
a  dog  and  then  making  a  fistulous  opening  between  the  skin  and  the  gall- 
bladder; all  the  bile  secreted  is  then  discharged  at  the  surface.  In  such 
animals  it  has  been  found  that  the  secretion  of  bile  is  continuous.  With 


FIG.  274. — Transverse  Section  through  Four  Crypts  of  Lieberkiihn,  from  the  Large 
Intestine  of  the  Pig.  They  are  lined  by  columnar  epithelial  cells,  the  nuclei  being  placed 
in  the  outer  part  of  the  cells.  The  divisions  between  the  cells  are  seen  as  lines  radiating 
from  L,  the  lumen  of  the  crypt;  G,  epithelial  cells,  which  have  become  transformed  into 
goblet  cells.  X  350.  (Klein  and  Noble  Smith.) 

the  great  discharge  of  bile  into  the  intestine  that  occurs  during  the  third 
hour  after  a  meal,  there  is  an  increased  secretion  of  this  fluid.  This  in- 
creased secretion  of  bile  can  also  be  evoked  by  the  introduction  of  o .  4  per 
cent,  hydrochloric  acid  into  the  duodenum,  and  occurs  even  after  the  di- 
vision of  all  connections  between  the  liver  and  the  central  nervous  system. 
There  is  evidence  that  the  increased  secretion  of  bile  is  brought  about 
through  a  mechanism  identical  with  that  for  the  secretion  of  pancreatic 
juice,  and  that  in  each  case  one  and  the  same  substance — secretin — is  formed 
by  the  action  of  the  cells  of  the  mucous  membrane  and  absorbed  into  the 
blood  stream  and  excites  both  the  liver  and  pancreas  to  increased  activity. 
The  emptying  of  the  gall-bladder  has  been  investigated  on  dogs  with  a 
Pavlov  fistula.  In  this  operation,  the  orifice  of  the  duct,  with  the  mucous 
membrane  around  it,  is  cut  out  of  the  wall  of  the  intestine  and  the  latter 


MODE    OF   SECRETION   AND   DISCHARGE    OF  BILE 


389 


again  closed.  The  excised  portion  with  the  opening  of  the  bile  duct  is 
stitched  into  the  abdominal  wound.  The  natural  orifice  of  the  duct  is  thus 
made  to  open  externally.  The  discharge  of  bile  is  found  to  begin  almost  im- 
mediately after  taking  food;  it  attains  its  maximum  during  tbe  third  hour, 
coincident  with  the  pancreatic  flow,  and  then  rapidly  diminishes.  Dale  has 
shown  that  the  muscular  fibers  of  the  wall  of  the  gall-bladder  are  supplied 
by  nerves  from  both  the  vagus  and  the  sympathetic.  The  former  are  motor, 
while  the  latter  convey  inhibitory  impulses.  The  contractions  of  the  gall- 
bladder are  provoked  reflexly  on  the  passage  of  the  acid  chyme  into  the 
intestine.  The  gall-bladder  acts  as  a  reservoir  for  the  bile  during  the  intervals 
when  digestion  is  not  in  progress.  The 
mechanism  by  which  the  bile  passes  into 
the  gall-bladder  is  simple.  The  orifice 
through  which  the  common  bile-duct  com- 
municates with  the  duodenum  is  narrower 
than  the  duct,  and  appears  to  be  closed, 
except  when  there  is  sufficient  pressure 
behind  to  force  the  bile  through  it.  The 
pressure  exercised  upon  the  bile  secreted 
during  the  intervals  between  periods  of 
digestion  appears  insufficient  to  overcome 
the  force  of  the  sphincter  by  which  the  orifice 
of  the  duct  is  closed;  and  the  bile  in  the 
common  duct  traverses  the  cystic  duct  and 
so  passes  into  the  gall-bladder.  It  is  proba- 
bly aided  in  this  retrograde  course  by  the 
peristaltic  action  of  the  ducts. 

The  bile   is   discharged  from  the  gall- 
bladder and  enters  the  duodenum  on  the 

introduction  of  food  into  the  small  intestine.  F!G-  ^.-Longitudinal.  Sec- 

lion  of  Fundus  of  Crypt  of  Lieber- 
It  is  pressed  on  by  the  contraction  of  the    kiihn.     b,    Goblet    cell    showing 

coats  of  the  gall-bladder  and  of  the  com-    mitosis;  ^epithelial  cell;  k  .cell  of 

Paneth;  /,  leukocyte  in  epithelium; 
mon  bile-duct.  m,  mitosis  in  epithelial  cell.     Sur- 

When  the  discharge  of  the  bile  into  the    rounding  the  crypt  is  seen  the 

stroma  of  the  mucous  membrane, 
intestine  is  prevented  by  an  obstruction  of     x  530.     (Kolliker.) 

some  kind,  as  by  a  gall-stone  blocking  the 

hepatic  duct,  it  is  reabsorbed  in  great  excess  into  the  blood,  and,  circu- 
lating with  it,  gives  rise  to  the  well-known  phenomena  of  jaundice.  This 
is  explained  by  the  fact  that  the  pressure  of  secretion  in  the  ducts, 
although  normally  very  low,  not  exceeding  15  millimeters  of  mercury 
in  the  dog,  is  still  higher  than  that  of  the  portal  veins.  If  the  pressure 
exceeds  15  mm.  the  secretion  continues  to  be  formed,  but  passes  into  the 
blood  vessels  through  the  lymphatics. 


3QO  FOOD   AND    DIGESTION 

The  Intestinal  Secretion,  or  Succus  Entericus.— It  is  impossible  to 
isolate  the  secretion  of  the  glands  of  Brunner  or  of  the  glands  of  Lieberkiihn, 
but  the  total  secretion  of  the  intestinal  mucosa  can  be  secured  by  isolating  a 
loop  of  intestine  by  the  operation  known  as  the  Thiry  fistula.  A  few  drops 
of  secretion,  the  succus  entericus,  can  be  obtained  by  this  means.  Intestinal 
juice  is  a  yellowish  alkaline  fluid  with  a  specific  gravity  of  i  .on  and  contains 
about  2  . 5  per  cent,  of  solid  matters. 

Intestinal  juice  has  only  slight  digestive  action.  It  contains  a  weak  pro- 
teolytic  enzyme  and  a  weak  amylolytic  enzyme.  Maltase  is  also  present. 
But  the  chief  and  most  profound  importance  is  given  to  the  intestinal  juice 
by  the  discovery  of  the  activating  enzyme,  enterokinase.  This  specific 
activating  enzyme  for  the  trypsinogen  of  the  pancreatic  juice  places  the  in- 
testinal secretion  in  the  rank  of  necessary  secretion  for  efficient  digestion. 
Enterokinase  can  be  prepared  by  extracting  the  superficial  scrapings  of 
the  intestinal  mucous  coat.  The  duodenal  region  is  richest  in  enterokinase, 
but  the  secretion  of  the  lower  intestinal  lengths  also  contains  the  enzyme. 

Extracts  of  the  mucosa  of  the  intestine  have  been  found  to  contain  an- 
other substance  which  has  the  specific  action  of  splitting  peptones  into 
simpler  polypeptids  and  amino-acids.  This  substance  has  been  called 
erepsin. 

There  are,  therefore,  three  important  new  substances  in  the  succus  en- 
tericus (or  in  the  extract  of  the  glands),  secretin  (page  360),  erepsin,  and 
enterokinase,  in  addition  to  the  proteolytic  and  diastatic  enzymes. 

Summary  of  the  Digestive  Changes  in  the  Small  Intestine. — The 
thin  chyme,  which,  during  the  whole  period  of  gastric  digestion,  is  being  con- 
stantly squeezed  or  strained  through  the  pyloric  orifice  into  the  duodenum, 
consists  of  albuminous  matter  that  is  breaking  down,  dissolving  and  half 
dissolved;  of  fatty  matter  that  is  mechanically  separated  and  melted,  but 
not  dissolved  at  all;  of  starch  in  various  stages  of  the  process  of  con- 
version into  sugar,  and  as  it  becomes  sugar  dissolving  in  the  fluids 
with  which  it  is  mixed;  and  with  these  are  mingled  gastric  juice  and 
fluid  that  has  been  swallowed,  together  with  such  portions  of  the  food  as 
are  not  digestible. 

The  chyme  in  the  duodenum  is  subjected  to  the  influence  of  the  bile  and 
pancreatic  juice  and  also  to  that  of  the  succus  entericus.  All  these  secretions 
have  a  more  or  less  alkaline  reaction,  and  at  once  neutralize  the  acid  of  the 
gastric  chyme. 

The  special  digestive  changes  in  the  small  intestine  are:  (i)  The  fats  are 
changed  by  the  bile  and  pancreatic  juice  in  two  ways:  (a)  They  are 
chemically  decomposed  by  the  alkaline  secretions,  and  a  soap  and  glycerin 
are  the  result.  (6)  They  are  emulsified;  i.e.,  their  particles  are  minutely 
subdivided  and  diffused,  so  that  the  mixture  assumes  the  condition  of  a 
milky  fluid  or  emulsion.  (2)  The  albuminous  substances  which  have  been 


ACTION  OF   MICRO-ORGANISMS   IN   THE    INTESTINES  391 

partly  dissolved  in  the  stomach  are  subjected  chiefly  to  the  action  of  the 
pancreatic  juice.  The  pepsin  is  rendered  inert  by  the  bile.  The  pancreatic 
trypsin  proceeds  with  the  further  conversion  of  the  proteoses  into  peptones, 
and,  with  the  erepsin,  of  the  peptones  into  leucin,  tyrosin,  and  the  other 
amino-acids.  (3)  The  starchy  portions  of  the  food  are  now  acted  on  briskly 
by  the  pancreatic  juice  and  the  succus  entericus,  and  are  changed  to  maltose 
and  dextrose.  (4)  Salines  are  usually  in  a  state  of  solution  before  they 
reach  the  intestine. 

Digestive  Changes  in  the  Large  Intestine. — The  changes  which  take 
place  in  the  chyme  in  the  large  intestine  are  probably  only  the  continuation 
of  the  same  changes  that  occur  in  the  course  of  the  food's  passage  through 
the  upper  part  of  the  intestinal  canal.  No  special  enzymes  have  been  clearly 
shown  for  the  mucous  membrane  of  the  large  intestine.  The  enzymes  of  the 
small  intestine  may  continue  their  action  here,  being  hindered  only  by  the 
acid  developed  from  fermentation  processes. 

Action  of  Micro-organisms  in  the  Intestines. — Certain  changes  take 
place  in  the  intestinal  contents  independent  of,  or  at  any  rate  supplemental 
to,  the  action  of  the  digestive  ferments.  These  changes  are  brought  about 


0°°. 

O 
"  O 


C  « 

I 


f 


FIG.  276. — Types  of  Micro-organisms,  a,  Micrococci  arranged  singly;  in  twos, 
diplococci — if  all  the  micrococci  at  a  were  grouped  together,  they  would  be  called  staphylo- 
cocci — and  in  fours,  sarcinae;  b,  micrococci  in  chains,  streptococci;  c,  and  d,  bacilli  of 
various  kinds,  one  is  represented  with  flagellum;  e,  various  forms  of  spirilla;/,  spores,  either 
free  or  in  bacilli. 

by  the  action  of  micro-organisms  or  bacteria.  We  have  indicated  elsewhere 
that  the  digestive  ferments  are  examples  of  unorganized  ferments,  so  bacteria 
are  examples  of  organized  ferments.  Organized  ferments,  of  which  the  yeast 
plant  may  be  taken  as  a  typical  example,  consist  of  unicellular  vegetable  organ- 
isms, which  when  introduced  into  a  suitable  medium  grow  with  remarkable 
rapidity.  By  their  growth  they  produce  new  substances  from  those  supplied  to 
them  as  food.  Thus,  for  example,  when  the  yeast  cell  is  introduced  intou  solu- 
tion of  grape-sugar,  it  grows,  and  alcohol  and  carbon  dioxide  are  produced. 
The  alcohol  and  carbon  dioxide  arise  from  the  formation  by  the  cell  of  some 
chemical  substances  which  are  allied  to  the  unorganized  ferments  and  which 
greatly  increase  in  amount  with  the  multiplication  of  the  original  cell.  In 


3Q2  FOOD   AND   DIGESTION 

all  such  fermentative  processes  organisms  analogous  to  the  yeast  cell  are 
present,  and  it  is  not  strange  that  if  the  ferment  cell  is  introduced  into  a 
suitable  medium  it  may  by  its  rapid  growth  convert  an  unlimited  amount  of 
one  substance  into  another.  Speaking  generally,  a  special  variety  of  cell 
is  concerned  with  each  ferment  action,  thus  one  variety  has  to  do  with 
alcoholic,  another  with  lactic,  and  another  with  acetous  fermentation. 

A  considerable  number  of  species  of  bacteria  exist  in  the  body  during  life 
chiefly  in  connection  with  the  mucous  membranes,  particularly  of  the  digest- 
ive tract.  Many  forms  of  bacteria  have  been  isolated  from  the  mouth,  a  few 
varieties  from  the  stomach,  and  a  very  large  number  from  the  intestines.  It 
is  only  in  the  last-named  locality  that  their  multiplication  has  much  effect 
from  a  physiological  point  of  view.  The  normal  (hydrochloric  acid)  acidity 
of  the  stomach  usually  destroys  all  the  micro-organisms  taken  in  with  the 
food,  but  when  the  amount  of  this  acid  is  deficient  (and  sometimes  even 
when  it  is  normal)  some  of  the  spores  may  escape.  On  reaching  the  small 
intestine  these  spores  begin  to  develop  in  its  alkaline  medium,  and  may  in- 
crease to  such  an  extent  as  to  stop  all  intestinal  digestion;  the  point  where 
this  occurs  varies  from  day  to  day.  The  large  intestine  always  swarms 
with  micro-organisms,  though  they  do  not  readily  pass  the  ileocecal  valve 
into  the  small  intestine.  Some  species  of  bacteria  found  in  the  intestine  are 
anaerobic;  i.e.,  they  do  not  develop  in  the  presence  of  free  oxygen. 

The  changes  induced  in  the  intestine  by  the  activity  of  micro-organisms 
are  of  two  kinds,  fermentation  and  putrefaction;  the  former  of  these  results 
in  the  breaking-down  of  carbohydrate  matter,  and  the  latter  in  the  disintegra- 
tion of  protein  matter.  The  process  of  fermentation  is  the  less  complex 
and  probably  occurs  normally  in  the  small  intestine.  The  lactic  acid  fermen- 
tation is  the  most  important,  though  the  butyric  acid  fermentation  is  next; 
under  the  influence  of  these  bacteria  the  carbohydrates  are  buty rates 
broken  down  into  lactic  and  butyric  acids,  and  perhaps  into  acetic  acid 
also.  Carbonic  acid  gas  may  be  formed  at  the  same  time  and  cause  flatu- 
lence. Cellulose  and  other  insoluble  carbohydrates  are  decomposed,  with 
the  formation  of  marsh  gas  and  hydrogen,  which  escape  by  the  rectum. 

In  putrefaction  the  process  is  somewhat  similar  to  that  in  tryptic  diges- 
tion, the  proteins  being  broken  down  into  peptones,  leucin,  tyrosin,  and  a 
long  row  of  similar  substances,  including  ammonia  in  relatively  large 
amount  from  the  large  intestine.  Folin  and  Denis  discovered  over  four 
times  as  much  ammonia  nitrogen  in  the  mesenteric  vein  as  in  the  carotid, 
0.38  to  0.44  milligrams  in  100  centimeters  of  the  blood  of  the  vein  and 
from  0.03  to  0.08  milligrams  in  the  same  amount  of  arterial  blood  from  the 
carotid.  It  also  results  in  the  production  of  various  gases,  such  as  carbon 
dioxide,  sulphureted  hydrogen,  ammonia,  hydrogen,  and  methane  (marsh 
gas),  and  of  a  high  percentage  of  the  volatile  fatty  acids,  valerianic  and 
butyric.  Of  the  aromatic  substances  the  most  important  are  some  phenol 
derivatives  and  indol  and  skatol,  though  their  toxicity  has  been  greatly 


THE    FECES 


393 


overestimated.  Indol  and  skatol  undergo  oxidation  in  the  liver  after  ab- 
sorption, forming  indoxyl  and  skatoxyl.  They  are  in  part  carried  off  in  the 
feces,  but  when  the  bowel  is  obstructed  they  are  absorbed  and  eventually 
appear  in  the  urine,  indoxyl  and  skatoxyl  forming,  respectively,  indoxyl- 
and  skatoxyl-sulphuric  acids  and  their  salts.  Tyrosin  is  broken  down  into 
para-oxy-phenol-propionic  acid,  paracresol,  and  phenol;  para-oxy-phenol- 
acetic  acid  is  also  formed.  The  phenols,  after  absorption,  are  in  part  con- 
jugated with  glycuronic  acid  which  is  formed  by  the  incomplete  oxidation 
of  dextrose  and  are  eliminated  into  the  urine.  Experiments  have  been  per- 
formed to  determine  whether  or  not  the  intestinal  bacteria  are  necessary  to 
normal  digestion.  The  weight  of  evidence  is  in  favor  of  the  view  that  they 
are  not. 

The  Feces. — The  contents  of  the  large  intestine,  as  they  proceed  toward 
the  rectum,  become  more  and  more  solid,  lose  more  liquid  and  nutrient 
parts,  and  gradually  acquire  the  odor  and  consistency  characteristic  of 
feces.  After  a  sojourn  of  uncertain  duration  in  the  sigmoid  flexure  of  the 
colon,  or  in  the  rectum,  they  are  finally  expelled  by  the  act  of  opening  the 
bowels,  or  defecation.  The  average  quantity  of  matter  evacuated  by  the 
human  adult  in  twenty-four  hours  is  about  200  to  250  grams,  but  the  amount 
and  character  vary  exceedingly  according  to  the  food  eaten.  Vegetable  foods 
contain  much  indigestible  matter,  while  meats  and  meat  diets  leave  very 
little  unabsorbed  material  to  be  expelled  in  the  feces. 

TABLE  OF  COMPOSITION  OF  FECES. 

The  amount  of  water  varies  considerably,  from  68  to  82  per  cent,  and 
upward.  The  following  table  gives  about  an  average  composition: 

Water 733 

Solids,  comprising: 

a.  Insoluble  residues  of  the  food,  uncooked  starch,  cellulose, 

woody  fibers,  cartilage,  horny  matter,  mucin,  seldom 
muscular  fibers  and  other  proteins,  fat,  and  cholesterin . 

b.  Certain  substances  resulting  from  decomposition  of  foods, 

such  as  indol,  skatol,  fatty  and  other  acids ;  calcium  and 
magnesium  soaps 

c.  Special  excretion, — excretin,  excretoleic  acid  (Marcet), 

and  stercorin  (Austin  Flint) 

d.  Salts — chiefly  phosphate  of  magnesium  and  phosphate  [-267 

of  calcium,  with  small  quantities  of  iron,  soda,  lime,  and 
silica 

e.  Insoluble  substances  accidentally  introduced  with  the 

food 

f.  Mucus,  epithelium,  altered  coloring  matter  of  bile,  fatty 

acids,  etc 

g.  Varying  quantities  of  other  constituents  of  bile  and  the 

secretions 

1000 


394 


FOOD   AND   DIGESTION 


Intestinal  Gases. — Under  ordinary  circumstances,  the  alimentary 
canal  contains  a  considerable  quantity  of  gases.  The  presence  of  gas  in  the 
intestines  is  so  constant  and  the  amount  in  health  so  uniform  that  there  can 
be  no  doubt  that  its  existence  is  a  normal  condition. 

The  gas  contained  in  the  stomach  and  bowels  is  from  air  swallowed  with 
either  food  or  saliva,  gases  developed  by  the  decomposition  of  foods,  or  of  the 
secretions  and  excretions  thrown  into  the  intestines.  The  decomposition  of 
foods  is  the  chief  source.  The  following  table,  compiled  by  Brinton,  is  a  col- 
lection of  analyses  that  have  been  made  and  is  chiefly  valuable  as  showing  the 
kinds  of  gases  present,  although  the  amounts  of  the  gases  vary  with 
the  diet. 

GASES  FOUND  IN  THE  ALIMENTARY  CANAL. 


Whence  obtained 

Composition  by  volume 

Oxygen 

Nitrog. 

Carbon 
dioxide 

Hydrogen 

Carburet, 
hydrogen 

Sulphuret. 
hydrogen 

Stomach  

II 

7' 
32 
67 

35 
46 

22 

14 
3° 
12 

51 
43 
40 

4 

38 
8 
6 

Small  intestines  .  . 

•  Trace. 
0-5 

Cecum  .  . 

I3 
8 
II 
19 

Colon  

Rectum   

Expelled  per  anum  



19 

An  analysis  of  the  intestinal  gases  by  Ruge  in  man  is  as  follows: 


Gases 

Milk  diet 

Meat  diet 

Vegetable  diet 

Carbon  dioxide  

o       to  1  6 

8      to  n 

2  I          to    7  A. 

Hydrogen  .  . 

A-t               +  O      tJJ. 

o   i  to     i 

Carbureted  hydrogen  

*to          **  DT- 
o  .  o 

u  .  y  vu      ^ 

2  6         to   3  7 

j.  .  5  MJ     4 

Ad.         to    ?  < 

Nitrogen  

•26      to  38 

A  f       to  6j. 

IO          to    IQ 

MOVEMENTS  .OF  THE  INTESTINES. 

The  muscular  activity  of  the  intestines  accomplishes  two  important  func- 
tions; i.e.,  it  thoroughly  mixes  the  digesting  food  and  secretions  and  it  carries 
the  content  along  the  tract.  Intestinal  peristalses  have  been  described  for  a 
long  time.  These  peristalses  begin  as  contractions  of  the  circular  muscles, 
producing  ring-like  constrictions  that  are  propagated  as  waves  over  the  intes- 
tine from  above  downward.  Such  constrictions  carry  the  intestinal  contents 
forward.  The  longitudinal  muscles  by  their  contraction  produce  pendular 
motion  of  the  intestine. 

A  most  instructive  contribution  to  the  knowledge  of  intestinal  movements 


MOVEMENTS    OF    THE    INTESTINES  395 

has  been  made  by  Cannon.  He  fed  cats  food  mixed  with  10  to  33  per  cent,  of 
subnitrate  of  bismuth,  and  observed  the  shadows  of  the  food  when  subjected 
to  the  X-rays.  A  length  of  food  in  the  intestine  was  seen  to  be  constricted 
into  a  series  of  oval  masses,  figure  277.  Each  of  these  oval  masses  is  quickly 
constricted  in  the  middle,  and  neighboring  halves  of  adjacent  masses  flow 
together.  After  this  process  is  repeated  a  number  of  times  a  peristaltic  wave 
of  the  type  previously  described  sweeps  the  whole  content  of  the  loop  down 
the  intestinal  tract.  The  segmentations  of  the  intestine  are  facilitated  by 
pressure  from  within,  perhaps  stimulated  by  the  direct  pressure  of  the  food. 
In  general,  hollow  organs  receive  their  most  effective  motor  stimulus  in  this 
manner,  i.e.,  by  distention.  This  rule  applies  in  the  lengths  of  the  small 
intestine. 


FIG.  277. — Diagram  Illustrating  the  Segmentation  of  the  Food  in  the  Small  Intestine. 

(Cannon.) 

Peristaltic  contractions  of  the  same  general  type  as  in  the  small  intestine 
also  occur  in  the  large  intestine.  Cannon  has  noted  a  variation  here  also. 
The  ascending  and  the  transverse  loops  of  the  colon  sometimes  exhibit 
rhythmic  antiperistalses  which  keep  the  content  moving  against  the  ilio- 
cecal  sphincter  for  several  minutes  at  a  time.  In  the  meantime  digesting 
material  is  being  received  into  the  ascending  colon  through  the  ilio-cecal 
sphincter,  being  slowly  forced  on  by  the  peristalses  of  the  lower  loops  of  the 
small  intestine.  From  time  to  time  strong  general  peristalses  in  the  colon 
slowly  force  the  food  onward.  Under  ordinary  circumstances  remnants 
of  unabsorbed  material  in  the  colon  do  not  pass  beyond  the  pelvic  colon,  be- 
ing held  at  this  point  until  the  greater  portion  of  the  entire  colon  is  moderately 
filled.  When  sufficient  material  has  accumulated  here,  it  is  evacuated  by 
strong  peristalses  combined  with  compression  by  the  contracting  abdominal 
muscles. 

Reverse  peristalsis,  antiperistalsis,  does  not  commonly  occur  in  the  small 
intestine,  but  large  nutrient  enemata  introduced  into  the  rectum  and  colon 
may  be  forced  by  antiperistaltic  waves  in  the  large  intestine  against  the 
ilio-cecal  sphincter  at  a  time  when  it  is  atonic  and  relaxed  thus  allowing 
materials  to  pass  through  the  ilio-cecal  valve  into  the  small  intestine.  In 
normal  healthy  individuals  this  sphincter  reacts,  according  to  Hertz,  like 
the  sphincter  of  the  pylorus  effectively  closing  the  tube. 


396  FOOD   AND    DIGESTION 

Influence  of  the  Nervous  System  on  Intestinal  Peristalsis. — As  in 

the  case  of  the  esophagus  and  stomach,  the  peristaltic  movements  of  the  in- 
testines may  be  directly  set  up  in  the  muscular  fibers  by  the  presence  of  food 
acting  as  the  stimulus.  Few  or  no  movements  occur  when  the  intestines  are 
empty,  vigorous  contractions  when  filled.  The  intestines  are  connected 
with  the  central  nervous  system  both  by  the  vagi  and  by  the  splanchnic  nerves, 
as  well  as  by  other  branches  of  the  sympathetic  which  come  to  them  from  the 
celiac  and  other  abdominal  plexuses.  The  relations  of  these  nerves  re- 
spectively to  the  movements  of  the  intestine  and  the  secretions  are  probably 
the  same  as  in  the  case  of  the  stomach  already  considered. 

The  vagus  carries  the  motor  fibers  for  the  intestine,  while  the  sympathetics 
are  inhibitory.  Various  states  of  the  central  nervous  system,  such  as  fear, 
anger,  etc.,  inhibit  the  intestinal  movements.  The  intestine,  and  stomach, 
too,  carries  out  peristalses  when  isolated  from  the  central  nervous  sys- 
tem and  indeed  from  the  body,  so  that  the  central  connections  do  not  originate 
the  rhythmic  stimulus  but  are  only  regulative.  The  intestinal  movements 
are  essentially  automatic.  It  has  long  been  known  that  isolated  portions 
of  the  intestine  contract  rhythmically  and  automatically.  This  has  been 
proven  for  both  the  longitudinal  and  circular  muscles,  and  in  the  absence 
of  the  mucous  membrane.  According  to  Magnus,  rhythmic  contractions 
do  not  occur  in  the  muscle  if  the  plexus  of  Auerbach  be  removed.  He  con- 
cludes that  the  automatic  rhythm  is  inherent  in  the  local  nerve  ganglia, 
that  it  is  not  a  reflex  since  it  occurs  in  the  absence  of  the  mucosa,  and  that 
the  contractions  of  the  smooth  muscles  are  directly  dependent  upon  the  local 
nerve  distribution.  Others  contend  that  the  contractions  are  independent, 
depending  on  the  rhythmic  property  of  the  muscle  itself,  but  co-ordinated 
by  the  complex  local  nervous  mechanism.  By  either  conception  the  nerve 
connections  with  the  central  nervous  system  are  regulative  and  co-ordinative. 

The  innervation  of  the  large  intestine  is  also  double  in  character  and 
the  relations  are  doubtless  the  same  as  in  the  small  intestine. 

Defecation. — The  emptying  of  the  bowels  is  essentially  an  involuntary 
act  which  has  acquired  a  certain  amount  of  voluntary  regulation.  The 
act  is  accomplished  wholly  reflexly  in  dogs  with  isolated  lumbar  cord,  in 
fact  has  been  observed  when  the  lumbar  spinal  cord  was  removed.  In 
the  latter  case  defecation  occurs  by  automatic  peristalsis  of  the  rectum,  and 
colon,  while  in  the  former  reflexes  through  the  lumbar  cord  carry  out  the 
act.  When  the  material  that  has  accumulated  in  the  colon  descends  into 
the  rectum,  which  is  normally  empty,  it  initiates  the  reflex  stimulus  which 
culminates  in  opening  the  bowel,  or  defecation.  Hertz  and  others  have 
taken  X-ray  photographs  of  the  human  immediately  before  and  after  open- 
ing the  bowels,  with  a  view  to  a  better  understanding  of  what  structures 
take  part  in  the  process.  It  is  found  that  the  entire  colon  as  far  back  as  the 
ilio-cecal  sphincter  may  be  emptied  during  the  act,  see  figure  2760. 


DEFECATION 


397 


EXAMPLE  2 


Normally  in  man  the  rectal  stimulus  gives  rise  to  the  consciousness  of 
the  desire  to  defecate  and  to  the  initiation  of  efferent  nerve  impulses  that  may 
increase  the  contraction  of  the  external  sphincter  and  inhibit  the  act  tempo- 
rarily. During  opening  of  the  bowels  however,  the  voluntary  effort  leads  to 
relaxation  of  the  external  sphincter,  and  the  normal  peristalsis  of  the  rectum 
is  suppported  by  contractions  of  the  abdominal  musculature  so  as  greatly 
to  increase  the  abominal  pressure,  thus  aiding  the  involuntary  reflex  which 
controls  the  relaxation  of 

the  internal  sphincter  and  EXAMPLE  i 

the    contractions    of    the 
colon,  and  rectum. 

The  Time  of  Passage 
of  Food  Through  the 
Alimentary  Canal. — It 
is  important  to  under- 
stand the  rapidity  with 
which  food  passes  along 
the  different  divisions  of 
the  alimentary  tube. 

The  effectiveness  of 
digestion,  the  dangers 
from  fermentation  and 
putrefaction,  and  the  ills 
that  follow  the  absorption 
of  toxic  by-products  are 
largely  dependent  upon 
the  time  element. 

At  the  beginning  of  a 
meal  the  activity  of  the 
entire  alimentary  canal  is 

low,  and  one  may  assume  that  the  various  muscular  mechanisms  are  re- 
latively relaxed.  When  food  is  swallowed,  immediately  small  portions 
begin  to  pass  through  the  stomach  and  pyloric  sphincter.  However,  the 
main  portion  of  the  food  passes  out  of  the  stomach  as  it  undergoes  digestion 
and  solution  during  the  succeeding  4  or  5  hours.  Hertz,  who  has  given  us 
the  most  accurate  picture  of  the  elements  in  this  process  allows  4^  hours 
for  the  passage  of  food  along  the  22  feet  of  the  small  intestine  to  the  iliac 
sphincter.  This  he  has  determined  by  X-ray  photographs  after  a  bismuth 
meal.  This  is  a  comparatively  rapid  passage  averaging  about  2\  cm.  per 
minute.  The  rate  is  much  slower  in  the  large  intestine.  He  allows  an 
average  speed  of  2  hours  each  for  the  ascending,  transverse,  and  descending 
colon.  In  the  short  lengths  of  the  iliac  and  pelvic  colons  an  equal  or  even 
greater  time  is  consumed.  Thus  he  would  allow  an  average  of  eighteen 


Immediately  after 


FIG.  2j6a. — The  upper  pair  of  figures  show  the 
bismuth  shadows  for  normal  defecation,  the  lower 
after  the  use  of  magnesium  sulphate.  (From  Hertz, 
Cook,  and  Schlesinger.) 


39$  FOOD   AND   DIGESTION 

hours  for  the  first  appearance  of  the  food  remnants  at  the  end  of  the  pelvic 
colon  where  it  is  retained  until  the  bowels  are  opened.  Of  course,  assuming 
an  empty  canal  ahead  of  the  moving  material,  the  undigested  and  waste 
products  of  the  food  require  on  an  average  from  eighteen  to  twenty-two 
hours  for  the  passage  of  the  entire  alimentary  tract.  The  earliest  possible 
moment  at  which  food  remnants  can  appear  in  the  feces  will  be  when  de- 
fecation occurs  just  after  those  remnants  which  passed  immediately  through 
the  stomach  have  reached  the  ascending  colon,  say  five  hours.  On  the  other 
hand,  assuming  that  a  meal  is  taken  on  an  entirely  empty  canal,  the  food 
remnants  will  not  produce  the  reflex  for  opening  the  bowels  until  they  reach 
the  pelvic  colon.  Hence  if  the  bowels  are  open  only  once  in  twenty-four 
hours,  it  is  obvious  that  the  remnants  of  a  meal  may  not  be  passed  before 
thirty-six  to  thirty-eight  hours. 


SALIVA    AND    SALIVARY    DIGESTION  399 

LABORATORY  EXPERIMENTS  IN  DIGESTION. 

I.  SALIVA  AND  SALIVARY  DIGESTION. 

1.  Reflex  Salivary  Secretion. — Saliva,  which  is  the  mixed  secretion 
of  the  salivary  and  buccal  glands,  is  produced  more  or  less  intermittently. 
Examine,  taste,  or  smell  appetizing  food,  for  example,  an  apple,  the  salivary 
glands  begin  to  discharge  secretion  which  is  poured  into  the  mouth  more 
rapidly  than  under  ordinary  conditions.     This  increased  activity  is  a  reflex 
secretion.     It  is  brought  about  by  the  stimulation  of  sensory  structures  which 
lead  to  afferent  nerve  impulses  reacting  on  nerve  centers  in  the  medulla  to 
cause  secretory  nerve  impulses  to  the  glands.     The  stimulating  effect  of  food 
in  the  mouth  causes  the  most  rapid  reflex  secretion,  which  may  last  through 
several   minutes   or   even   hours.     Especially   stimulating   substances   are, 
beside  food,  such  substances  as  tartaric  acidj  lemon  juice,  ether,  alcohol,  etc., 
in  fact,  anything  that  produces  strong  local  irritation  will  lead  to  reflex 
secretion. 

2.  The  Secretory  Nerves  of  the  Salivary  Glands  of  the  Dog. — The 
nervous  mechanism  for  the  salivary  glands  is  well  known,  and  the  anato- 
mical relations  are  such  as  to  make  these  glands  favorable  for  studying  the 
nervous  mechanism  of  glands  in  general. 

Anesthetize  a  dog  and  bind  it  to  a  suitable  holder.  Expose  the  nerves  to 
the  submaxillary  gland  as  follows:  cut  through  the  skin  of  the  lower  jaw 
along  the  inner  border  for  about  3  inches.  Isolate  and  double  ligate  the 
jugular  vein  and  any  other  veins  in  the  field  except  the  ones  coming  from  the 
submaxillary  gland.  Isolate  and  cut  the  digastric  muscle,  also  the  mylo- 
hyoid,  using  pains  not  to  injure  the  duct  of  the  gland  or  its  arteries.  When 
the  muscles  are  laid  back',  the  artery  and  accompanying  sympathetic  nerve 
branches,  the  hypoglossal  and  the  lingual  nerves,  the  submaxillary  duct  and 
the  submaxillary  gland,  will  all  be  exposed.  Isolate  and  introduce  a  very 
fine  glass  cannula  into  the  submaxillary  duct  tying  it  firmly  in  place.  A  small 
nerve  filament  branches  from  the  lingual  nerve  and  runs  to  the  hilus  of  the 
gland,  the  chorda  tympani.  Carefully  expose  the  chorda,  place  a  silk  liga- 
ture under  it  for  convenience  in  handling.  Also  expose  the  sympathetic 
filaments  with  the  artery. 

Stimulate  the  chorda  tympani  with  a  mild  induction  current  for  a  few 
1  minutes  at  a  time  at  intervals,  and  note  that  the  secretion  which  is  absent  or 
forming  slowly  before  stimulation  now  gathers  quickly  and  leaves  the  end 
of  the  cannula  in  a  series  of  drops.  Collect  the  saliva  in  a  small  beaker.  One 
can  measure  the  rate  of  flow  by  collecting  the  saliva  in  a  small  graduated 
cylinder  or,  by  changing  the  beaker  every  ten  minutes,  making  a  record  of 
the  quantity  of  secretion  formed.  Stimulate  the  sympathetic  fibers,  cutting 
the  hypoglossal  nerve  if  necessary,  and  note  that  the  secretion  is  very  slightly 


400  FOOD    AND    DIGESTION 

increased,  but  the  increase  lasts  for  only  a  few  minutes.  If  the  sympathetic 
fibers  are  stimulated  before  the  chorda,  then  the  sympathetic  secretion  is 
relatively  less  than  if  the  order  of  stimulation  is  reversed. 

Connect  the  duct  with  a  mercury  manometer  and  record  the  secretion 
pressure,  compare  with  arterial  blood  pressure.  It  is  often  greater.  Explain. 

During  stimulation  of  the  nerves,  note  the  relative  flow  of  blood  through 
the  organ.  During  chorda  stimulation  the  flow  is  increased;  during  sympa- 
thetic stimulation  it  is  decreased,  as  these  nerves  contain  vaso-dilator  and 
vaso-constrictor  fibers,  respectively. 

Inject  5  milligrams  of  pilocarpine.  This  drug  stimulates  the  secre- 
tory nerve  endings  and  thus  causes  a  copious  flow  of  saliva. 

3.  Microscopic  Changes  in  the  Gland  Cells. — Make  a  histological 
preparation  (by  any  standard  method  of  fixing  and  staining)  of  the  submaxil- 
lary  gland  of  the  cat,  a,  taken  after  a  period  of  several  hours'  fasting  when 
the  gland  cells  may  be  assumed  to  be  at  rest;  and  b,  immediately  after  a  period 
of  activity  (from  eating,  or  activity  secured  by  the  stimulation  of  the  chorda 
tympani)  and  note :  a,  The  cells  from  the  resting  gland  are  relatively  larger, 
the  nuclei  are  pushed  back  against  the  basement  membrane,  they  have 
sparsely  sustaining  protoplasm,  and  the  cells  are  crowded  with  large  gran- 
ules, which  in  a  fortunate  preparation  fill  the  entire  cell.     The  outlines  of  the 
cells  are  relatively  indistinct  and  the  lumen  of  the  gland  is  small,     b,  The 
cells  of  the  active  gland  are  relatively  small,  the  nuclei  are  centrally  placed, 
the  protoplasm  stains  more  definitely,  the  granules  are  usually  present  but 
limited  to  the  side  of  the  cell  next  to  the  lumen,  the  outlines  of  the  cells  are 
distinct,  and  the  lumen  is  often  quite  large. 

4.  The  Chemical  Composition  of  Saliva. — Collect  several  cubic  cen- 
timeters of  saliva  as  follows:     Wash  the  mouth  thoroughly  with  water,  then 
induce  secretion  of  saliva  by  chewing  a  bit  of  paraffin  or  a  piece  of  thoroughly 
washed  rubber.     The  inhalation  of  ether  vapor  will  often  facilitate  the  reflex 
secretion.     One  should  avoid  strong  acids  to  induce  secretion  unless  their 
presence  is  to  be  taken  into  consideration  afterward.     Make  the  following 
tests: 

Reaction. — A  slip  of  neutral  litmus-paper  when  introduced  into  freshly 
collected  saliva,  or  for  convenience  simply  held  in  the  mouth  during  sali- 
vary secretion,  shows  an  alkaline  reaction. 

Mucin. — To  3  or  4  cc.  of  saliva  add  2  per  cent,  acetic  acid  drop  by  drop 
until  distinct  acidity  is  obtained.  On  stirring  the  saliva  with  a  glass  rod  a 
sticky  mucin  makes  its  appearance. 

Potassium  Sulphocyanide. — To  2  cc.  of  saliva  in  a  test-tube  add  2  or  3 
drops  of  ferric  chloride  solution,  slightly  acidulated  with  hydrochloric  acid, 
a  reddish-brown  coloration  indicates  the  presence  of  potassium  sulpho- 
cyanide.  One  should  run  a  blank  test  on  distilled  water  for  comparison. 


INFLUENCE    OF    ACIDS    AND    ALKALIES    ON    SALIVARY   DIGESTION    401 

Chlorides. — Add  silver  nitrate  to  2  cc.  of  saliva  after  first  removing 
the  proteins.  A  white,  cloudy  precipitate,  which  disappears  on  adding 
ammonia  and  reappears  on  adding  nitric  acid,  indicates  the  presence  of 
chlorides. 

Proteins. — Remove  the  mucin  from  a  sample  of  saliva,  as  above,  and  test 
by  the  characteristic  protein  reactions.  A  faint  trace  of  protein  can  usually 
be  demonstrated. 

5.  Digestive  Action  of  Saliva  on  Starch. — Review  the  tests  for  starch, 
dextrin,  and  dextrose,  as  preparation  for  an  identification  of  these  products 
of  salivary  digestion.     To  10  cc.  of  i  per  cent,  starch  paste  in  the  water-bath 
at  40°  C.,  add  2  cc.  of  saliva,  and  mix  thoroughly  with  a  glass  rod.     Immedi- 
ately begin  two  series  of  tests:  a,  for  the  presence  of  starch;  b,  for  the  presence 
of  reducing  sugar.     The  tests  for  starch  can  be  made  by  adding  to  3  drops 
of  solution,  on  a  porcelain  plate,  an  equal  quantity  of  dilute  iodine  in  potas- 
sium iodide  solution.     Use  a  glass  rod.     Make  the  tests  every  2  minutes 
for  20  minutes.     The  tests  for  reducing  sugar  are  best  made  by  placing  2  cc. 
of  Fehling's  solution  in  each  of  a  series  of  test-tubes  and  adding  to  suc- 
cessive tubes,  at  intervals  of  5  minutes,  i  cc.  portions  of  the  digest  from  a 
dropping  pipet  and  boiling.     If  the  tests  are  set  away  as  fast  as  they  are  pre- 
pared, a  reddish-yellow  cuprous  oxide  will  settle  out,  and  the  series  will  give 
a  rough  comparison  as  to  the  quantity  of  reducing  sugar  present. 

In  the  first  series  the  deep  blue  of  the  starch  reaction  quickly  changes 
to  a  reddish-blue,  a  red,  a  reddish-brown,  until  finally  no  change  in  color 
other  than  that  produced  by  the  mixture  of  the  iodine  occurs,  showing  that 
the  starch  has  passed  the  second  stage  of  erythro-dextrin  in  its  disappearance. 
The  indication  of  reducing  sugar  in  the  second  series  shows  that  this  erythro- 
dextrin  has  been  transformed  into  reducing  sugar,  and  also  that  the  amount 
of  sugar  is  greatly  increased  during  the  progress  of  the  test. 

6.  The  Influence  of  Temperature  on  Salivary  Digestion. — Prepare 
three  test-tubes,  a,  6,  c,  containing  i  cc.  each  of  saliva.     Boil  a,  place  b  in  a 
water-bath  at  40°  C.,  and  place  c  in  ice  water.     After  c  has  been  cooled  down 
to  the  temperature  of  the  ice-bath,  add  to  each  2  cc.  of  i  per  cent,  starch  solu- 
tion and  mix.     At  intervals  of  5  minutes  test  these  three  samples  for  the 
disappearance  of  starch  and  appearance  of  reducing  sugar,  as  in  Experi- 
ment 5.     No  change  will  take  place  in  a;  b  will  be  quickly  digested,  and  the 
digestion  in  c  will  be  slight  or  suspended.     Upon  placing  c  in  a  warm  bath 
digestion  will  quickly  occur. 

7.  Influence  of  Acids  and  Alkalies  on  Salivary  Digestion. — To 
each  of  5  test-tubes,  a,  b,  c,  d,  e,  add  2  cc.  saliva  and  water,  or  the  solutions 
given  in  the  table,  so  that  the  saliva  will  be  uniformly  diluted.     Then  add 
quickly  i  cc.  of  i  per  cent,  starch  paste  to  each.     Run  parallel  tests  for  the 

appearance  of  reducing  sugar  and  disappearance  of  starch. 

26 


402 


FOOD    AND    DIGESTION 


The  results  obtained  in  Experiments  5,  6,  and  7  show  that  starch  is 
converted  into  reducing  sugar,  and  furthermore,  that  the  conditions  for  its 
conversion  indicate  that  the  change  is  accomplished  by  an  amylolytic  enzyme 
which  in  this  case  is  called  ptyalin. 


Steps 

A                       B 

C 

D 

E 

i.  Prepare  and 
set  in  water- 
bath  at  40°  C. 

2  cc.  saliva 
2  cc.   water 

2  cc.  saliva 
2  cc.  0.5  per 
cent,     sodium 
bicarbonate 

2  cc.  saliva 

2    CC. 

strong 
sodium 
hydrate 

2  cc.  saliva 
2  cc.  0.4  per 
cent,  hydro- 
chloric acid 

2  cc.  saliva 
2  cc.  strong  hy- 
drochloric acid 

2.  Then  add 

i  cc.  starch 

i  cc.  starch 

i  cc.  starch 

i  cc.  starch 

i  cc.  starch 

3.  Test  for 
starch  and 
sugar 
immediately 

4.  After  10 
minutes 

5.  After  20 
minutes 

6.  After  40 
minutes 

8.  The  Action  of  Ptyalin  is  Favored  by  the  Removal  of  the  End 
Products. — Place  50  cc.  of  2  per  cent,  starch  paste  in  a  dialyzing  tube  or 
paper,  suspend  in  a  beaker  of  running  water.  Take  50  cc.  of  the  same 
solution  in  a  beaker,  to  each  add  2  cc.  of  saliva  and  mix  thoroughly.  Test 
for  the  disappearance  of  starch  at  intervals  of  20  minutes.  The  starch  in 
the  dialyzing  tube  will  disappear  first  because  the  reducing  sugar  passes  out 
through  the  dialyzer,  while  in  the  beaker  it  is  retained  and  hinders  the  further 
action  of  ptyalin. 


II.  GASTRIC  JUICE  AND  GASTRIC  DIGESTION. 

9.  The  Secretion  of  Gastric  Juice. — The  conditions  which  influence 
gastric  secretion  can  be  readily  observed  on  the  dog  with  a  gastric  fistula. 
Take  a  dog  which  has  had  a  gastric  fistula  prepared  some  weeks  before  and 
which  is  in  a  condition  of  hunger,  place  him  in  a  holder  with  a  cup  suspended 
to  collect  the  gastric  juice,  and  exhibit  before  the  dog  some  fresh  meat  or 
other  food  which  he  enjoys,  but  do  not  allow  him  to  eat  it.  After  teasing  the 


COMPOSITION    OF    GASTRIC    JUICE 


403 


FIG.  278. — Operation  on  the  Stomach  to  Form 
an  Isolated  Pouch  with  Nerves  Intact.  S, 
Isolated  sac;  V,  cavity  of  stomach;  A,  A,  open- 
ing at  the  abdominal  wall. 


animal  for  5  to  10  minutes,  an  abundant  flow  of  gastric  juice  will  begin. 
Pawlow  calls  this  the  psychic  secretion. 

If  an  esophageal  fistula  has  also  been  performed  the  dog  may  be  allowed 
to  eat  the  meat,  of  course  swal- 
lowing it  out  of  the  esophageal 
fistula  back  into  the  plate.  In 
this  experiment  an  abundant 
flow  of  gastric  secretion  takes 
place  and  may  continue  for  an 
hour  or  more. 

If  a  gastric  pouch  has  been 
performed  by  Pavlov's  method, 
the  animal  may  be  allowed  to 
eat  the  food,  swallowing  it  into 
the  stomach.  In  this  case  the 
reflex  secretion  just  described 
takes  place  as  usual,  but  is  fol- 
lowed after  an  hour  or  an  hour 
and  a  half  by  a  second  increase 
in  the  quantity  of  secretion. 
This  second  increase  has  been 
ascribed  to  the  reflexes  origi- 
nating in  the  stomach,  possibly  from  the  reflex  stimulating  action  of  the 
digestive  products  themselves. 

10.  Composition  of  Gastric  Juice. — Test  a  sample  of  gastric  juice 
obtained  from  a  gastric  fistula,  as  follows: 

Reaction. — Gastric  juice  is  strongly  acid.  Test  for  free  hydrochloric  acid 
as  follows:  Gastric  juice  turns  congo-red  to  a  blue  color.  Gastric  juice  plus 
0.5  per  cent,  alcoholic  solution  of  dimethyl-amido-azobenzol  develops  a 
cherry-red  color,  a  reaction  that  is  given  by  free  hydrochloric  acid.  Com- 
bined hydrochloric  acids  and  organic  acids  do  not  give  the  color.  Giinz- 
burg's  reagent,  consisting  of  2  per  cent,  phloroglucin  and  i  per  cent,  vanillin 
in  80  per  cent,  alcohol,  produces  a  rose-colored  mirror  on  porcelain  in  the 
presence  of  free  hydrochloric  acid.  The  test  is  very  delicate. 

The  lactic  acid  sometimes  present  in  the  contents  of  the  stomach  is 
derived  partly  from  the  sarcolactic  acid  of  muscle  and  partly  from  lactic-acid 
fermentation  of  carbohydrates.  Lactic  acid  (C3HSO3),  if  present,  gives  the 
following  test:  A  solution  of  10  cubic  centimeters  of  a  4  per  cent,  aqueous 
solution  of  carbolic  acid,  20  cubic  centimeters  of  water,  and  one  drop  of  ferric 
chloride  is  made;  forming  a  blue-colored  mixture.  A  mere  trace  of  free  lactic 
acid  added  to  such  a  solution  causes  it  to  become  yellow.  Inasmuch  as 
mineral  acids  also  discharge  the  color,  the  lactic  acid  should  first  be  removed 
from  the  gastric  contents  by  shaking  with  ether  and  the  test  tried  out  with  a 
solution  of  the  residue  after  evaporation  of  the  ether. 


404  FOOD    AND    DIGESTION 

Proteins. — The  usual  protein  tests  (page  107)  can  be  applied  to  gastric 
juice  and  show  that  it  contains  small  quantities. 

11.  Artificial  Gastric  Juice. — The  active  principle,  pepsin,  of  gastric 
juice  can  be  obtained  by  extracting  the  gastric  mucous  membrane  of  the 
dog,  pig,  etc.     Scrape  off  the  mucous  membrane,  grind  it  to  a  fine  pulp  by 
repeatedly  running  it  through  a  sausage  machine  or  by  pounding  in  a  mortar 
with  clean  sand.     The  mucous  membrane  should  be  allowed  to  stand  for 
three  or  four  hours  before  extraction,  otherwise  the  zymogen,  and  not  the 
enzyme,  will  be  obtained.    Extract  a  portion  of  this  gastric  pulp  in  water  and 
filter.     Or  extract  with  glycerin  for  several  weeks  and  filter.     Either  of  these 
extracts  contains  the  enzyme.     A  solution  of  the  glycerin  extract  in  o .  2  per 
cent,  hydrochloric  acid  contains  all  the  properties  of  gastric  juice.     This 
solution  is  known  as  artificial  gastric  juice. 

Commercial  pepsin  already  prepared  can  be  obtained  on  the  market. 
Artificial  gastric  juice  is  made  from  commercial  pepsin  by  adding  0.5 
grams  scale  pepsin  per  hundred  cc.  of  0.2  per  cent,  hydrochloric  acid,  which 
gives  a  very  active  preparation. 

12.  Digestive  Action  of  Gastric  Juice,  or  Artificial  Gastric  Juice 
on  Proteins. — The  chief  digestive  action  of  gastric  juice  is  on  proteins. 
Shreds  of  fibrin  which  permit  the  gastric  juice  to  come  in  intimate  contact 
with  all  parts  of  the  material  form  the  best  protein  for  testing  the  action  of 
this  enzyme.     Prepare  a  series  of  test-tubes,  a,  b,  c,  d,  each  containing  5  cc. 
of  artificial  gastric  juice.     Add  to  a  some  shreds  of  fibrin;  to  b  some  boiled 
white  of  egg;  to  c  some  fibers  of  boiled  meat;  to  d  some  fibers  of  raw 
meat;  place  in  a  warm  bath  at  40°  C.  and  examine  at  intervals  of  20 
minutes.     Tabulate  the  results  by  the  plan  indicated  in  Experiment  13, 
noting  particularly  the  rapidity  with  which  the  different  proteins  go  into 
solution. 

13.  Condition  Affecting  the  Enzyme  Action  of  Gastric  Juice. — Pre- 
pare a  series  of  test-tubes  containing  5  cc.  each  of  gastric  juice,  according 
to  the  table  on  the  next  page.     Add  fibrin  threads  to  each  and  note  the 
changes  at  intervals  of  10  minutes. 

14.  The  Effect  of  Bile  on  Peptic  Digestion. — The  influence  of  bile 
on  the  activity  of  pepsin-hydrochloric  acid  is  demonstrated  in  the  following 
steps: — i.  Place  in  a  series  of  test  tubes,  A,  B,  C,  D,  2  cc.  each  of  0.5  per 
cent,  pepsin  hydrochloric  acid  solution.     2.  Add  to  A    2  cc.  of  water;  B  2 
cc.  water,  o.i  cc.  bile  solution  (5  per  cent,  dried  bile);  C  1.5  cc.  water,  0.5 
cc.  bile  solution;  D  2  cc.  bile  solution.     3.  Add  to  each  test  tube  2  cc.  of 
0.4  per  cent,  hydrochloric  acid  and  shake  thoroughly.     Drop  in  each  test 
tube  4  fibrin  threads,  and  set  in  water  bath  at  40°  C.     Tabulate  the  results 
at  intervals  of  10  minutes. 

15.  Cleavage  Products  of  Gastric  Digestion. — Add  5  to  10  grams 
of  fibrin  to  500  cc.  of  artificial  gastric  juice  in  a  flask  and  place  in  a  water- 
bath  at  40°  C.     After  one  hour  filter  off   100  cc.     Exactly   neutralize  this 


CLEAVAGE    PRODUCTS    OF    GASTRIC    DIGESTION 


405 


filtrate  with  i  per  cent,  potassium  hydrate.  A  precipitate  makes  its  appear- 
ance, and  can  be  collected  on  the  filter-paper,  washed  with  distilled  water, 
and  dissolved  in  i  per  cent,  hydrochloric  acid,  acid  metaprotein.  Test 
for  the  protein  reactions. 


A 

B 

C 

D 

E 

F 

2  cc.  pepsin 

2cc.  hydro- 

2cc. pepsin 

2cc.  pepsin 

2cc.  pepsin 

2cc.  pepsin- 

solution  .5 

chloric  acid 

hydro- 

hydro- 

hydro- 

alkali solu- 

Prepare 

per  cent,  in 

0.4  per  cent. 

chloric 

chloric 

chloric 

tion.  (0.5  per 

water 

acid  solu- 

acid solu- 

acid solu- 

cent. 

(neutral) 

tion,  (arti- 

tion in  ice 

tion, 

NaHC03) 

ficial  gas- 

bath 

boiled 

tric  juice) 

Then  add 

4  fibrin 

4  fibrin 

4  fibrin 

4  fibrin 

4  fibrin 

4  fibrin 

threads 

threads 

threads 

threads 

threads 

threads 

Changes 

after  10 

minutes 

after  20 

minutes 

after  40 

minutes 

after  60 

minutes 

After  twelve  hours  or  more  filter  the  remaining  400  cc.,  exactly  neutralize 
to  remove  any  traces  of  acid  albumin,  and  filter.  The  filtrate  contains  pro- 
teoses  and  peptones.  Concentrate  the  filtrate  over  a  water-bath  to  one- 
fourth  its  volume,  add  an  equal  quantity  of  saturated  ammonium  sulphate 
solution,  a  sticky  precipitate  of  primary  proteases  separates  out.  Collect  on 
a  filter-paper  or  in  a  centrifuge,  wash  with  half-saturated  ammonium 
sulphate,  redissolved  in  very  dilute  salt  solution,  and  test  for  protein 
reactions.  The  primary  proteoses  are  precipitated  by  nitric  acid. 

To  the  filtrate  from  the  half-saturated  ammonium  sulphate  add  crystals 
of  ammonium  sulphate  until  complete  saturation  with  the  salt.  Deutero- 
albumoses  separate  out.  Collect  on  a  filter-paper,  wash,  dissolve,  and  test 
for  proteins.  The  secondary  proteoses  are  not  precipitated  by  nitric  acid. 

Finally  the  filtrate  contains  peptone.  It  can  be  isolated  and  tested  by 
concentrating  over  the  water-bath,  adding  barium  hydrate  to  slight  excess 


4C>6  FOOD   AND    DIGESTION 

to  remove  the  sulphate,  filtering,  and  precipitating  the  excess  of  barium  by 
exact  neutralization  with  i  per  cent,  sulphuric  acid.  Test  for  protein  reac- 
tions. Peptone  gives  a  rose  color  in  the  biuret  reaction.  The  xanthoproteic 
reaction  gives  the  color  change,  but  not  the  usual  precipitate.  Peptone  is  re- 
dissolved  from  its  alcoholic  precipitate  without  change.  It  is  dialyzable. 

1 6.  Action  of  Rennin. — Add  a  solution  of  commercial  rennin  (jun- 
ket powder),  or  of  the  extract  of  gastric  mucous  membrane  of  the  fourth 
stomach  of  the  calf,  to  5  cc.  of  milk  and  let  stand  for  a  few  minutes.     Repeat 
the  test  with  artificial  gastric  juice.     Also  with  neutral  gastric  juice.     In  each 
case  tne  milk  will  form  a  jelly-like  clot,  which  is  firmer  in  the  test-tube  con- 
taining commercial  rennin.     In  the  test-tube  containing  artificial  gastric 
juice  the  milk  is  first  coagulated,  then  slowly  dissolved  or  digested.     This 
clotting  is  due  to  the  special  coagulating  enzyme,  rennin. 

III.  PANCREATIC  JUICE  AND  PANCREATIC  DIGESTION. 

17.  The  Secretion  of  Pancreatic  Juice. — If  a  dog  containing  a  pan- 
creatic fistula  made  by  Pavlov's  method  is  available,  then  try  the  experi- 
ment of  feeding  the  animal  and  noting  the  rate  of  secretion  of  pancreatic 
juice  through  a  period  of  two  hours.     When  the  gastric  digestion  has  pro- 
ceeded to  the  point  where  the  acid  chyme  may  be  supposed  to  have  entered 
the  duodenum,  then  a  sharp  increase  in  the  flow  of  pancreatic  juice  takes 
place.     This  increased  activity  will  last  through  a  period  of  two  or  three 
hours  or  more.    It  is  produced  either  by  nerve  reflexes  (Pavlov)  or  by  the 
influence  of  the  secretin  produced  by  the  intestinal  mucous  membrane  when 
stimulated  by  acid. 

18.  Influence  of  Secretin   on  the  Rate   of  Secretion. — Make  an 
extract  of  the  intestinal  mucous  membrane  from  the  duodenum,  by  scrap- 
ing off  the  membrane,  grinding  it  to  a  pulp,  and  extracting  it  over  a  water- 
bath  in  0,2  per  cent,  hydrochloric  acid,  and  filtering. 

Anesthetize  a  large  dog,  open  the  abdomen,  isolate  the  pancreatic  duct, 
introduce  a  cannula,  and  arrange  for  the  collection  of  pancreatic  juice.  Intro- 
duce a  cannula  into  the  saphenous  vein  and  connect  it  with  a  buret  containing 
the  extract  of  secretin  already  prepared.  Inject  5-cc.  quantities  of  the 
secretin  solution  into  the  vein  at  intervals  of  ten  minutes.  Measure  the  rate 
of  secretion  of  pancreatic  juice  by  counting  the  drops  per  minute  or,  if  the 
secretion  is  rapid  enough,  by  collecting  it  at  intervals  of  five  or  ten  minutes 
and  measuring  it  in  a  graduated  pipet. 

This  method  will  often  yield  enough  pancreatic  juice  in  the  course  of  a 
couple  of  hours  to  make  the  pancreatic  experiments  which  follow.  Bayless 
and  Starling  call  it  secretin  juice. 

19.  Chemical  Characters  of  Pancreatic  Juice. — Test  the  reaction, 
protein,  salt,  etc.,  content  of  the  sample  of  pancreatic  juice  collected  in  the 
last  experiment. 


CLEAVAGE    PRODUCTS    OF    PANCREATIC    DIGESTION  407 

20.  Artificial   Pancreatic   Juice. — Artificial   pancreatic  juice  can  be 
prepared  from  the  pancreas  by  grinding  and  macerating  and  extracting  a 
pancreas  with  water  or  glycerin,  as  described  for  the  gastric  glands  in  Experi- 
ment ii  above.     Commerical  preparations  of  pancreatic  enzyme  can  be 
obtained  on  the  market.     A  solution  of  a  glycerin  extract  of  pancreatic  gland 
or  of  commercial  pancreatin  in  o .  2  per  cent,  sodium  carbonate  is  known  as 
artificial  pancreatic  juice. 

21.  The  Enzymes  of  Pancreatic  Juice. — The  pancreatic  juice  con- 
tains enzymes  which  exert  a  digestive  action  on  starches,  fats,  and  proteins. 
This  fact  can  be  tested  as  follows:  a,  to  5  cc.  of  artificial  pancreatic  juice 
add  2  cc.  of  i  per  cent,  starch  paste,  mix  and  set  in  the  water-bath  at  40°  C.; 

b,  to  i  cc.  of  pancreatic  juice  (artificial  juice  is  not  active),  collected  in  Ex- 
periment 17,  add  0.5  cc.  of  neutral  olive  oil,  and  place  over  a  water-bath; 

c,  to  5  cc.  of  artficial  pancreatic  juice  add  a  few  flocks  of  fibrin.     Test  the 
digestive  action  on  starch  by  the  iodine  test  for  the  disappearance  of  starch, 
or  by  the  copper-reduction  test  for  the  presence  of  reducing  sugar.     Test  the 
fat  by  its  reaction,  noting  that  the  neutral  or  slightly  alkaline  solution  has 
become  acid,  also  by  the  fact  that  an  emulsion  has  been  formed.     Note  that 
the  protein  has  gone  into  solution. 

The  digestive  action  on  starch  is  due  to  the  enzyme  amylopsin,  or  pan- 
creatic diastase,  as  it  is  sometimes  called.  The  fat-splitting  effect  is  due  to  the 
enzyme  lipase,  and  the  solution  of  the  fibrin  is  accomplished  by  the  proteolytic 
enzyme,  trypsin. 

22.  Conditions  which  Influence   the  Action   of  the  Enzymes   of 
Pancreatic  Juice. — Prepare  each  of  5  test-tubes,  a,  b,  c,  d,  e,  as  shown  in 
the  table  on  the  next  page.     Place  a,  b,  c,  d  in  the  water-bath  at  40°  C. ,  and 
e  into  an  ice-bath.      Add  to  each  tube  2  cc.  of  i  per  cent,  starch  paste.     Follow 
the  digestive  changes  by  the  tests  previously  outlined  and  tabulate  the  results. 

Repeat  this  experiment  with  a  second  set  of  test-tubes  containing  4 
threads  of  fibrin  in  each.  Lipase  is  not  very  active  in  artificial  pancreatic 
juice  and  may  be  omitted,  but  if  pancreatic  juice  is  available  make  a  third 
set  containing  fat. 

23.  Cleavage  Products  of  Pancreatic  Digestion. — To  400  cc.  of  arti- 
ficial pancreatic  juice  add  25  grams  of  moist  fibrin  and  place  in  a  water-bath 
at  40°  C.,  add  2  cc.  of    chloroform  or  of  thymolin  alcoholic  solution  to 
prevent  putrefactive  changes.     After  three  or  four  hours  filter  off  100  cc. 
and  place  the  remainder  on  the  water-bath  for  one  or  two  days.     Test  the 
filtrate  for  alkali  ablumin,  albumoses,  and  peptones  by  the  method  outlined 
in  Experiment  14  above. 

Filter  the  second  portion  and  concentrate  to  a  syrupy  mass  on  the  water- 
bath.  Crystals  make  their  appearance.  Pour  off  the  fluid,  wash  the  crystals 
with  cold  water,  and  examine  under  the  microscope  for  sheaves  of  tyrosin. 
The  filtrate  contains  leucin. 


408 


FOOD   AND    DIGESTION 


If  the  digestion  had  been  allowed  to  proceed  without  the  antiseptic, 
bacteria  would  have  appeared  in  the  solution,  and  protein  cleavage  products, 
due  to  their  action,  would  be  found,  notably  indol,  phenols,  and  volatile 
fatty  acids. 


A 

B 

C 

D 

E 

Take 

2  cc.  neutral 
pancreatin, 
2  cc.   0.5   per 
cent.  Na2CO3 

2  cc.  neutral 
pancreatin, 
2  cc.    water 

2  cc.  neutral 
pancreatin, 
2  cc.  0.4  per 
cent.  HC1 

2  cc.  neutral 
pancreatin, 
2  cc.  water. 
Boil 

2  cc.  neutral 
pancreatin, 
2  cc.  Na2CO3. 
Keep  at  o°  C. 

Then  add 

2  cc.  of  starch 

2  cc.  of  starch 

2  cc.  of  starch 

2  cc.  of  starch 

2  cc.  of  starch 

Note  after 
20   min- 
utes 

After     40 
minutes 

After     60 
minutes 

IV.  BILE  AND  INTESTINAL  JUICE. 

24.  Bile. — Secure  bile  from  the  gall-bladder  of  a  pig  or  dog,  or,  if 
it  is  possible,  a  sample  of  human  bile.     Test  the  reaction  which,  in  fresh 
bile,  is  neutral.     Test  for  mucin,  albumin,  and  for  iron;  hydrochloric  acid 
and  ferrocyanide  of  potassium  give  a  blue  color  when  iron  is  present. 

BileSalts. — Evaporate  10  c.c.  of  bile  to  complete  dryness,  mix  with  animal 
charcoal,  add  50  c.c.  of  absolute  alcohol,  filter;  add  an  excess  of  ether  to  the 
nitrate,  which  gives  a  white  precipitate  of  bile  salts.  Crystals  will  form  on 
standing  in  a  well -stoppered  flask  for  a  day  or  two. 

Bile  Acids. — A  drop  of  syrup  of  cane-sugar  in  a  test-tube  of  bile  forms  a 
deep  red-purple  color  at  the  line  of  separation  from  concentrated  sulphuric 
acid.  Furfur  aldehyde  with  cholalic  acid  gives  the  color. 

Bile  Pigments. — With  i  c.c.  of  bile  in  a  test-tube  strong  nitroso-nitric 
acid  produces  a  play  of  colors  beginning  with  green,  blue,  red,  and  yellow— 
Gmelin's  test. 

Bile  does  not  contain  digestive  enzymes,  but  the  bile  wets  the  mucous 
surface  of  the  intestine  and  facilitates  the  solution  and  absorption  of  fat  and 
fatty  acids. 

25.  Intestinal  Juice. — The  secretion  of  the  mucous  membrane  of  the 
small  intestine  has  been   proven  to  have  a  weak  digestive  action  on  pro- 
teins and  perhaps  on  starches.     It  can  be  obtained  from  an  intestinal  fistula. 


ALIMENTARY    MECHANISMS  409 

Its  chief  digestive  importance  consists  in  the  presence  of  the  activating 
enzyme,  enterokinase.  Enterokinase  can  be  prepared  by  extracting  the 
mucous  membrane  of  the  small  intestine  by  the  method  outlined  for  making  a 
pancreatic  extract. 

To  two  test-tubes  containing  2  cc.  of  artificial  pancreatic  juice,  or  pref- 
erably containing  secretin  pancreatic  juice,  add  2  threads  of  fibrin.  Keep  one 
for  the  control,  to  the  other  add  2  cc.  of  enterokinase  solution.  The  test- 
tube  containing  enterokinase  will  digest  more  rapidly  and  more  effectively 
than  the  other. 

V.  ALIMENTARY  MECHANISMS. 

26.  Normal  Peristalsis  and  the  Vagus  Control  of  the  Frog  Stomach 
and  Intestine. — Pin  a  pithed  frog  supine  on  a  frog  board.     Expose  the 
stomach  and  isolate  the  vagus.     Attach  a  lever  to  the  outer  curvature  of 
the  stomach  by  S-shaped  hook  and  thread,  so  that  the  contractions  register 
as  usual.     Anchor  the  gastric  mesentery  by  pin  but  avoid  blood  vessels. 

a.  Make  a  continuous  record  on  a  slow  drum  of  the  normal  peristalsis 
of  the  stomach  for  30  minutes. 

b.  Stimulate  the  vagus  nerve  for  5  to  10  seconds  with  weak  induction 
shocks,  as  tested  by  your  tongue.     Mark  the  time  of  stimulation  by  a 
signal  magnet.     Allow  a  long  recovery  period  after  each  stimulation. 
Repeat  with  stronger  stimuli. 

c.  Compare  by  direct  inspection  the  general  characteristics  of  gastric 
peristalsis  both  before  and  following  vagus  stimulation.     Does  stimulation 
induce  a  single  contraction  or  a  rhythm? 

d.  Transfer  the  lever  to  the  duodenum  and  record  its  normal  con- 
tractions.    Then   stimulate    the   vagus    to    test   extrinsic   motor   nerve 
control. 

e.  Expose  the  small  intestine  of  the  frog  just  used  and  make  obser- 
vations by  inspection  without  registration.     Lightly  pinch  the  pyloric 
stomach  or  duodenum  with  a  forceps.     Note  the  time  with  watch  and 
determine  the  rate  of  peristalsis.     Observe  the  final  swinging  movements 
of   the  cloaca  or  rectum.     Remove  the  intestine  and  measure  its  total 
length.     Calculate  the  rate  of  propagation  per  centimeter  per  second  and 
the  total  time  for  the  wave  to  pass  the  length  of  the  intestine. 

27.  Gastro-intestinal  Movements  of  the  Cat. — Use  a  two  kilo  cat 
which  has  been  fed  ground  meat  four  hours  before.     Give  10  cc.  of  a  10  per 
cent,  chloral  hydrate  solution  per  rectum,  or  4  grams  of  urethane,  twenty 
minutes  before  the  experiment.     Very  little  ether  is  then  required  for 
narcosis.     Open  the  abdomen  along  the  midline  to  expose  the  stomach  and 
intestine.     Protect  the  visceral  organs  by  covering  with  very  thin  wax  paper. 


410  FOOD   AND    DIGESTION 

a.  Observe  two  kinds  of  intestinal  movements,  segmenting  movements 
of   the  intestine,   and  peristaltic  movements,  i.e.,  rings  of  contractions 
moving  downward.     Compare   these   movements   carefully,   noting   the 
rate  of  propagation,  the  sequence,  etc. 

b.  Note  especially   the   movements  of  the  stomach.     Compare   the 
pylorus  with  the  fundus.     Where  does  peristalsis  commence  in  the  cat 
stomach? 

c.  Insert  a  stomach  tube  and  fill  with  warm  water  until  distended, 
record  any  reactions. 

d.  Stimulate  the  vagus  in  the  neck  with  interrupted  induction  shocks 
of  moderate  intensity. 

e.  Cut  the  splanchnics,  then  stimulate  the  vagus  again.     Explain  the 
variations. 

/.  Inject  i  cc.  of  a  o.i  per  cent,  nicotine  solution  intravenously, 
explain. 

g.  Inject  i  cc.  of  o.i  per  cent,  epinephrin. 

28.  Pancreatic  and  Bile  Secretion. — Anesthetize  and  tracheotomize 
a  dog  and  connect  with  the  ether  apparatus.  Expose  the  jugular  for 
venous  injections.  Ligate  the  vagus  on  the  same  side  and  section. 
Open  the  abdomen  along  the  upper  half  of  the  linea  alba,  6  or  8  inches. 
Expose  the  duodenum  and  insert  a  cannula  into  the  common  bile  duct 
at  the  point  where  it  joins  the  intestine.  The  greater  pancreatic  duct 
along  the  posterior  border  of  the  pancreas  enters  the  intestine  below  the 
bile  duct.  Carefully  dissect  and  insert  a  cannula.  Provide  all  the  can- 
nulae  with  small  rubber  tubes  long  enough  to  extend  to  the  surface  through 
the  wound.  Carefully  adjust  the  parts  and  close  the  wound  by  stitches. 

a.  Record  the  rate  of  pancreatic  and  bile  secretions  by  signal  key. 

b.  Stimulate  the  peripheral  end  of  the  vagus  in  the  neck.     Give  time 
for  recovery  from  the  vascular  effects  and  for  the  secretion  to  develop. 

c.  Inject  20  to  30  cc.  of  0.4  per  cent,  hydrochloric  acid  by  hypodermic 
needle  into  the  duodenum.     The  reaction  is  slow.     Allow  15  minutes  or 
more  and  repeat. 

d.  Inject  40  cc.  of  "secretin"  solution  prepared  by  extracting  the 
macerated    mucosa    of    the    duodenum   with   0.4   per    cent.    HC1   and 
neutralizing    carefully    and    filtering.     Secretin    is    in    solution    in    the 
filtrate.     Repeat  in  20  to  30  minutes  (Journal  of  Physiology,  Vol.   28, 
P-  235>  1920).     Read  the  topic  of  chemical  control  and  hormones. 


CHAPTER  IX 
ABSORPTION 

THE  term  absorption  in  its  restricted  physiological  use  means  the 
process  by  which  the  digested  foods  pass  through  the  walls  of  the  alimentary 
canal  and  into  the  circulation.  In  a  more  general  sense  absorption  is  the 
process  by  which  substances  pass  from  one  part  of  the  body  to  another  by 
means  other  than  the  blood  and  lymph  vessels.  Usually  absorption  takes 
place  from  a  free  surface,  such  as  the  mucosa  of  the  alimentary  canal,  the 
surface  of  the  skin,  and  from  the  lungs. 

The  alimentary  canal  is  lined  throughout  with  a  continuous  layer  of  epi- 
thelial tissue.  This  layer  is  only  a  single  cell  thick  in  most  of  its  extent, 
but  nevertheless  it  effectively  separates  the  food  inside  the  canal  from  the 
lymph  in  the  tissue  interspaces  on  the  outside  of  the  mucous  membrane. 
These  spaces  are  separated  from  the  blood  in  the  adjacent  blood  vessels  by  a 
second  continuous  layer,  the  endothelial  walls  of  the  capillaries.  The  food, 
therefore,  during  its  absorption  from  the  alimentary  canal  must  pass  through 
two  layers  of  tissue  to  reach  the  blood  stream.  But  the  submucous  lym- 
phatic spaces  and  vessels  furnish  channels  which  may  carry  substances  into 
the  blood  by  way  of  the  thoracic  duct.  The  mucous  membrane  is,  therefore, 
the  one  strict  barrier  through  which  the  food  must  pass  in  the  act  of  absorption. 

The  exact  methods  by  which  absorption  takes  place  have  long  been  a 
subject  of  controversy  and  of  research.  But  this  problem  is  of  such  diffi- 
culty that  it  is  yet,  in  the  main,  unsolved.  Known  physical  and  chemical 
laws  are  thought  to  explain  the  facts  of  absorption.  Some  of  the  phys- 
ical factors  concerned  in  absorption  and  elimination  have  already  been 
considered  in  a  former  chapter,  osmosis  and  diffusion,  Chapter  IV.  A  third 
factor,  filtration,  consists  in  the  passage  of  a  fluid  under  pressure  through  a 
membrane.  These  factors  undoubtedly  play  an  important  role  in  the  pas- 
sage of  solutions  through  the  alimentary  mucous  membrane  and  the  walls  of 
the  blood  vessels.  The  part  which  the  physical  factors  play  is  probably  more 
pronounced  in  the  absorption  of  water  and  crystalloids.  The  nature  of  the 
fluid  within  the  digestive  tract,  and  the  movements  of  the  walls  of  the  stomach 
and  intestines  by  means  of  which  the  material  to  be  absorbed  is  brought 
into  intimate  contact  with  the  absorbing  membrane,  are  additional  factors 
which  influence  absorption. 

But  the  mechanical  and  physical  factors  do  not  fully  explain  the  observed 
facts  of  absorption.  It  becomes  more  and  more  evident  that  there  is  an 
unexplained  factor  bound  up  in  the  characteristics  of  the  living  protoplasm 
of  the  epithelial  cells  themselves.  When  isotonic  blood  serum  is  introduced 

411 


412  ABSORPTION 

into  the  intestine  the  salts  and  water  are  at  once  absorbed,  also  the  albumins, 
but  more  slowly.  In  this  experiment  the  osmotic  conditions  are  in  balance 
and  the  pressure  is  greater  on  the  side  of  the  blood  vessels,  so  that  absorption 
takes  place  with  the  actual  expenditure  of  energy.  The  important  fact 
here  is  that  the  absorption  through  a  living  membrane  is  influenced  by  the 
membrane  in  ways  that  we  cannot  yet  explain.  It  is  this  factor  which  de- 
termines the  different  rate  of  absorption  and  the  so-called  selective  absorp- 
tion in  different  regions  of  the  alimentary  canal. 

As  a  rule,  the  current  of  absorption  is  from  the  stomach  or  intestine  into 
the  blood;  but  the  reversed  action  may  occur,  as,  for  example,  when  sulphate 
of  magnesium  is  taken  into  the  alimentary  canal.  In  this  case  there  is  a 
rapid  discharge  of  water  from  the  blood  vessels  into  the  canal.  The  rapidity 
with  which  matters  may  be  absorbed  and  diffused  through  the  textures  of 
the  body  has  been  found  by  experiment.  It  appears  that  lithium  chloride 
may  be  diffused  into  all  the  vascular  textures  of  the  body,  and  into  some 
of  the  non-vascular,  as  the  cartilage  of  the  hip-joint,  as  well  as  into  the  aque- 
ous humor  of  the  eye,  in  a  quarter  of  an  hour  after  being  given  by  way  of  the 
mouth  and  on  an  empty  stomach.  Lithium  carbonate,  when  taken  in  five- 
or  ten-grain  doses  on  an  empty  stomach,  may  be  detected  in  the  urine  in 
five  or  ten  minutes;  or,  if  the  stomach  be  full  at  the  time  of  taking  the  dose, 
in  twenty  minutes. 

Absorption  in  the  Mouth. — The  epithelial  lining  of  the  mouth  is 
of  the  thicker  stratified  type  and  the  conditions  are  otherwise  unfavorable 
for  absorption.  Little,  if  any,  absorption  normally  takes  place  in  the  mouth, 
and  the  same  is  true  for  the  esophagus. 

Absorption  in  the  Stomach. — The  mucous  and  submucous  coats  of 
the  stomach,  see  figure  258,  are  well  supplied  with  blood  vessels  and  lym- 
phatics. The  mucous  membrane  is,  however,  so  crowded  with  the  peptic 
glands  that  the  relative  amount  of  absorbing  surface  is  small.  It  is  limited 
to  the  mucous  membrane  around  the  mouths  of  the  glands. 

Recent  experiments  have  shown  that  though  absorption  does  take  place 
in  the  stomach,  it  is  not  as  active  as  was  formerly  supposed,  even  in  the  case 
of  water.  Von  Mering  has  found  that  water  begins  to  pass  from  the  stomach 
into  the  intestine  almost  as  soon  as  it  is  swallowed,  and  that  very  little  of  it 
is  absorbed  from  the  stomach.  Of  500  cc.  given  by  the  mouth  to  a  large 
dog  with  a  duodenal  fistula,  only  5  cc.  were  absorbed  in  25  minutes,  the 
rest  having  passed  into  the  intestine.  Peptones,  sugars,  and  salts  are  ab- 
sorbed in  the  stomach,  but  only  to  a  limited  extent.  Peptones  are  not  ab- 
sorbed in  appreciable  amount  unless  present  to  as  much  as  5  per  cent,  or 
more.  Examination  of  the  mucous  membrane  during  the  stage  of  active 
digestion  has  revealed  the  presence  of  albumoses.  Sugars,  like  peptones,  are 
absorbed  by  the  stomach  only  to  a  slight  extent  in  the  weaker  solutions, 
but  are  readily  absorbed  when  the  more  concentrated  solutions  are  intro- 


ABSORPTION    IN    THE    STOMACH 


413 


duced  into  the  stomach,  5  per  cent,  and  over  (von  Mehring).  That  fats  are 
absorbed  in  the  stomach  was  clearly  indicated  by  von  Kolliker  as  far  back  as 
1857,  although  the  fact  seems  to  have  been  more  or  less  ignored  all  these 
years.  He  observed  an  increase  in  the  amount  of  fat  in  the  gastric  mucosa 
of  both  young  and  old  animals  after  feeding.  This  observation  has  been 
confirmed  under  carefully  guarded  experimental  conditions,  not  only  for 
the  different  experimental  laboratory  animals,  but  for  snakes  and  a  number  of 
fishes.  The  relative  amount  of  fat  absorbed  through  the  gastric  mucosa  is 
small,  however,  compared  with  that  absorbed  by  the  intestinal  villi.  Even 


•: 


FIG.  279. — Scheme  of  Blood  Vessels  and  Lymphatics  of  Human  Small  Intestine,  a, 
Central  lacteal  of  villus;  b,  lacteal;  c,  stroma;  d,  muscularis  mucosa^;  e,  submucosa;/,  plexus 
of  lymph  vessels;  g,  circular  muscle  layer;  h,  plexus  of  lymph  vessels;  i,  longitudinal 
muscle  layer;  j,  serous  coat;  k,  vein;  /,  artery;  m,  base  of  villus;  n,  crypt;  o,  artery  of  villus; 
p,  vein  of  villus;  q,  epithelium.  (Mall.) 

salts  in  the  stomach  are  not  readily  absorbed  until  the  concentration  is  from 
three  to  four  times  that  of  the  blood.  This  fact  is  in  direct  opposition  to  the 
popular  views  on  the  subject. 

While  some  absorption  does  take  place  in  the  stomach  it  is  evidently  not 
of  any  great  importance  under  normal  conditions.  The  presence  of  alcohol 
has  been  shown  to  increase  the  amount  of  absorption,  and  pepper,  mustard, 
and  such  drugs  as  produce  mild  local  irritation  accomplish  the  same  result. 


414  ABSORPTION 

Absorption  in  the  Intestines. — The  products  of  digestion  are  all  readily 
absorbed  in  the  small  intestine,  as  is  abundantly  shown  by  experiments. 
Absorption  from  the  small  intestine  has  been  studied  in  the  human  subject 
in  the  case  of  a  patient  who  had  a  fistulous  opening  in  the  lower  part  of  the 
ileum.  For  example,  85  per  cent,  of  the  protein  of  a  test  meal  was  absorbed 
before  the  food  reached  the  fistula.  The  food  passes  slowly  down  the  length 
of  the  small  intestine,  and  the  digestive  changes  produce  a  series  of  cleavages 


Lymphatics  of  head  and     I      ~"':  -  "'ffittHlBi^fe^:;^  '- »  *V^afcSl    Lymphatics  of  head  and 
neck,  right  '  HlttyS&OT  •'  •&& •••-  xfKS^         neck,  left 

'  '•nl'QBBtHf^ScfuiK^ 

Right  internal  jugular  vein     j  S^UHHflK^^S    Thoracic  duct 

Right  subclavian  vein     .___ ___„„ ... ,„____. 

^m.*      ^K«**  -»«B««KB^«I^^B«HU*       Left  subclavian  vein 
Lymphatics  of  right  arm 


Thoracic  duct 


Receptaculum  chyli 

Lacteals 


Lymphatics  of  lower  ex-  SVV^UHi^BYT^^]  I    Lymphatics  of  lower  ex- 

tremities EnJKIvVHBHHlf'nU^avHP        tremities 

FIG.  280. — Diagram  of  the  Principal  Groups  of  Lymphatic  Vessels.     (From  Quain.) 

which  have  known  osmotic  and  diffusion  properties.  The  question  has  been 
to  determine  which  of  the  cleavage  products  are  most  favorable  for  absorp- 
tion and  the  details  of  the  mechanism. 

The  mucous  membrane  of  the  small  intestine  possesses  special  structures 
for  absorption,  the  villi.  Each  villus  projects  as  a  finger-like  process  into 
the  lumen  of  the  intestine.  Its  single-layered  covering  of  epithelial  cells 
supported  by  a  connective-tissue  framework  brings  a  relatively  large  extent 
of  surface  into  contact  with  the  digesting  food,  which  is  thus  separated  from 
a  loop  of  capillaries  and  lymphatic  radicals. 


ABSORPTION  IN  THE  INTESTINES 


415 


The  capillaries  of  the  villus  are  connected  with  the  veins  which  contribute 
to  the  portal  vein,  hence  carry  blood  to  the  liver.  The  lacteals  of  the  villus 
contribute  to  the  mesenteric  lacteal  system,  hence  the  chyle  and  lymph  pass 
through  the  mesenteric  glands  and  the  thoracic  duct  to  the  subclavian  vein 


FIG.  281. 


FIG.  282. 


FIG.  281. — Superficial  Lymphatics  of  the  Forearm  and  Palm  of  the  Hand,  ^. — 5,  Two 
small  glands  at  the  bend  of  the  arm;  6,  radial  lymphatic  vessels;  7,  ulnar  lymphatk  vessels; 
8,  8,  palmar  arch  of  lymphatics;  9,  9',  outer  and  inner  sets  of  vessels;  &,  cephalic  vein;  d, 
radial  vein;  e,  median  vein;/,  ulnar  vein.  The  lymphatics  are  represented  as  lying  on  the 
deep  fascia.  (Mascagni.) 

FIG.  282. — Lymphatic  Vessels  of  the  Head  and  Neck  and  the  Upper  Part  of  the  Trunk. 
(Mascagni.)  £. — The  chest  and  pericardium  have  been  opened  on  the  left  side,  and  the 
left  mamma  detached  and  thrown  outward  over  the  left  arm,  so  as  to  expose  a  great  part 
of  its  deep  surface.  The  principal  lymphatic  vessels  and  glands  are  shown  on  the  side 
of  the  head  and  face,  and  in  the  neck,  axilla,  and  mediastinum.  Between  the  left  internal 
jugular  vein  and  the  common  carotid  artery,  the  upper  ascending  part  of  the  thoracic  duct 
marked  i,  and  above  this,  and  descending  to  2,  the  arch  and  last  part  of  the  duct.  The 
termination  of  the  upper  lymphatics  of  the  diaphragm  in  the  mediastinal  glands,  as  well  as 
the  cardiac  and  the  deep  mammary  lymphatics,  is  also  shown. 


416 


ABSORPTION 


in  the  neck.  There  are  thus  two  routes  by  which  absorbed  foods  may  reach 
the  general  circulation.  These  paths  can  be  independently  isolated;  and  a 
study  of  the  composition  of  their  discharge  during  active  absorption  con- 
tributes to  our  knowledge  of  the  course  taken  by  the  different  absorption 
products. 

Absorption  of  Proteins  from  the  Intestines. — Protein  is  absorbed 
chiefly  in  the  small  intestine,  though  just  exactly  how  cannot  at  present  be 
affirmed.  In  the  preceding  chapter  the  cleavage  products  of  protein  diges- 
tion have  been  discussed.  It  was  shown  there  that  proteoses,  peptones, 


FIG.  283. — A  Small  Portion  of  Medullary  Substance  from  a  Mesenteric  Gland  of  the 
Ox.  d,  d,  Trabeculae;  a,  part  of  a  cord  of  glandular  substances  from  which  all  but  a  few  of 
the  lymph  corpuscles  have  been  washed  out  to  show  its  supporting  meshwork  of  retiform 
tissue  and  its  capillary  blood  vessels  (which  have  been  injected  and  are  dark  in  the  figure); 
b,  b,  lymph  sinus,  of  which  the  retiform  tissue  is  represented  only  at  c,  c.  X  300.  (Kolliker.) 

peptids,  and  the  amino-acids  are  derived  from  the  proteins  as  digestion 
products.  It  has,  in  the  past,  been  assumed  that  peptone  represents  the 
form  most  freely  absorbed.  No  peptone  has,  however,  been  isolated  from 
the  blood  or  lymph  on  the  vascular  side  of  the  epithelial  membrane.  The 
present  supposition  is  that  the  protein  cleavage  products  are  taken  up  by 
the  epithelium  and  synthesized  into  other  and  more  complex  forms  before 
being  discharged  into  the  blood;  or  that  they  are  resynthetized  into  the  charac 
teristic  tissue  proteins  after  absorption.  The  digestion  cleavages  not  so 
utilized  are  desamidized  by  the  liver,  and  the  ammonia  so  formed  subse- 
quently converted  into  and  eliminated  as  urea. 

In  animal  foods,  such  as  eggs,  meat,  etc.,  it  is  estimated  that  about  98 
per  cent,  of  the  protein  is  absorbed;  whereas  in  vegetable  foods,  where  the  pro- 


ABSORPTION    OF    CARBOHYDRATES   BY    THE    INTESTINES  417 

tein  is  often  protected  from  the  action  of  the  digestive  enzymes,  there  may 
be  10  to  15  per  cent.  loss.  Analysis  of  the  total  lymph  discharged  by  the 
thoracic  duct  fails  to  show  any  increase  of  proteins  during  active  digestion, 
from  which  it  is  inferred  that  proteins  pass  by  way  of  the  liver. 

The  non-nitrogenous  residue  is  oxidized  or  temporarily  stored  as  glycogen. 

From  12  to  15  per  cent,  of  the  protein  remains  in  the  food  as  it  passes  the 
ileocecal  valve.  This  amount,  less  the  loss  in  the  feces,  is  absorbed  in  the 
large  intestine. 

Absorption  of  Carbohydrates  by  the  Intestines.— Carbohydrates 
are  broken  down  to  dextrose,  levulose,  etc.,  and  are  absorbed  as  such.  Even 


FIG.  284. — Section  of  the  Villus  of  a  Rat  Killed  during  Fat  Absorption,  ep,  Epithelium; 
sir,  striated  border;  c,  lymph  cells;  c',  lymph  cells  in  the  epithelium;  /,  central  lacteal  con- 
taining disintegrating  lymph  corpuscles.  (E.  A.  Schafer.) 

the  soluble  cane-sugar  is  split  by  the  invertase  of  the  intestine  into  the  mono- 
saccharides,  dextrose  and  levulose.  Starch  is  the  source  of  most  of  the  500 
grams  of  dextrose  absorbed  in  an  average  diet  per  day.  During  the  absorp- 
tion of  a  carbohydrate  meal  the  percentage  of  dextrose  in  the  blood  of  the 
portal  vein  is  increased  over  the  normal,  which  is  o .  i  to  1.5  per  cent.  This 
excess  of  dextrose  passes  through  the  liver  and  is  temporarily  stored  in  the 
liver  cells  as  glycogen.  In  the  case  of  a  fistula  in  the  receptaculum  chyli. 
the  chyle  contained  less  than  a  half  per  cent,  of  the  total  dextrose  absorbed. 

Experiments  on  the  rate  of  absorption  of  the  different  sugars  seem  to 
indicate  that  their  absorption  does  not  follow  known  physical  laws  and  that 
we  must  assume  an  unknown  chemical  factor  in  the  living  protoplasm. 

Dextroses  are  absorbed  readily  by  the  large  intestine. 


4i8 


ABSORPTION 


FIG.  285. — Mucous  Membrane 
of  Frog's  Intestine  during  Fat  Ab- 
sorption, ep,  Epithelium;  str, 
striated  border;  C,  lymph  corpus- 
cles; /,  lacteal.  (E.  A.  Schafer.) 


Fermentation  processes  from  bacterial  growth  produce  certain  acids  from 
the  carbohydrates,  chiefly  in  the  large  intestine.  These  are  readily  absorbed. 
Absorption  of  Fats  by  the  Intestines.— Fats  reach  the  absorbing 
epithelium  in  two  forms,  as  soluble  glycerol  and  soaps  and  as  finely  emulsi- 
fied fats.  The  first  two  are  taken  up  by  the  epithelium  readily  enough, 
but  the  proof  of  absorption  of  emulsified  fats  is  not  so  clear.  It  is  compara- 
tively easy  to  demonstrate  the  presence  of  microscopic  globules  of  fat  in  the 
intestinal  mucosa  both  in  the  epithelial  cells  themselves  and  to  a  less  degree 

in  the  intercellular  substance.  But  it  has 
been  constantly  noticed  that  there  is  a  clear 
zone  along  the  inner  or  free  borders  of  the 
cells.  Fat  drops  exist  in  the  adjacent 
digesting  mass,  and  in  the  deeper  parts  of 
the  cells,  but  not  in  this  border  zone.  The 
demonstration  of  the  reversible  action  of 
lipase,  has  strengthened  Pfliiger's  fat  dis- 
sociation theory  which  holds  that  before 
absorption  the  emulsified  fats  too  must  be 
decomposed.  They  can  then  pass  through 
the  cell  border  and  are  resynthesized  in 
the  cell  protoplasm.  This  is  of  course 
against  the  strictly  mechanical  view,  which 

must  be  abandoned  in  the  presence  of  the  evidence  supporting  the  newer 
conception.  The  decrease  in  efficiency  of  fats  as  foods  when  the  bile,  which 
wets  the  mucous  surface  and  dissolves  the  fatty  acids,  is  withheld  from  the  in- 
testine, supports  this  view.  As  absorption  progresses  the  size  of  the  fat 
drops  in  the  epithelial  cells  increases,  a  fact  that  is  readily  explained  by  sup- 
posing a  continued  synthesis  and  accumulation  of  fat.  Pfliiger's  view 
of  absorption  has  recently  received  strong  support  in  the  observations  of 
Bloor  that  isomannid  esters  of  fatty  acids  when  fed  to  animals  were 
digested  but  could  not  be  recovered  after  absorption.  Supposing  that 
lipolytic  cleavage  occurred  in  this  fat  during  digestion,  it  would  of  course 
not  be  rebuilt  in  the  cells  of  the  epithelium  after  absorption.  On  the 
theory  that  the  absorption  of  fats  takes  place  in  the  emulsified  form,  this 
compound  should  have  reappeared  in  the  chyle,  but  it  did  not. 

The  fat  is  ultimately  discharged  into  the  connective-tissue  spaces,  passes 
through  the  lymph  channels,  the  thoracic  duct,  and  into  the  blood  of  the  sub- 
clavian  vein.  This  is  the  course  taken  by  the  larger  percentage  of  the  fat. 
However,  during  absorption  some  of  the  fat  enters  the  capillaries  of  the  villi 
and  passes  through  the  liver.  The  presence  of  fat  drops  in  the  liver  cells 
at  certain  times  can  be  ascribed  to  storage  of  this  absorbed  fat,  the  liver 
exhibiting  not  only  a  glycogenic  but  a  lipogenic  function. 

It  is  said  that  the  more  readily  emulsified  fats,  those  that  melt  readily  at 


ABSORPTION  OF  MINERALS  AND  WATER  IN  THE  INTESTINES     419 

the  body  temperature,  are  the  more  completely  absorbed.  The  efficiency 
of  such  absorption  is  as  high  as  96  to  98  per  cent,  for  the  oils,  and  decreases 
sharply  for  such  fats  as  the  tallows. 

The  large  intestine  is  capable  of  absorbing  fats,  though  not  so  readily  as 
the  small  intestine. 

Absorption  of  Minerals  and  Water  in  the  Intestines.— The  salts 
common  in  the  foods  are  most  of  them  readily  soluble,  dissociate  quite  com- 
pletely in  the  dilute  solutions,  and  diffuse  and  dialyze  readily.  Of  the  salts 
of  the  foods,  the  sodium  and  potassium  cations  and  chlorine  anion  are  the 
most  readily  dissociated  and  are  most  diffusible,  while  the  calcium  and 
magnesium  cations  and  the  sulphate  anion  are  least  diffusible.  The  sub- 
stances pass  through  the  intestinal  epithelial  cells  and  the  intercellular  sub- 
stance; at  least  salts  easily  recognized  by  microchemical  means  have  been 
found  in  both  localities  during  absorption.  It  seems  probable  that  the  forces 
concerned  are  largely  osmosis  and  diffusion. 

Yet  observers  have  not  been  able  to  show  that  the  rate  and  character  of 
the  absorption  of  even  the  salines  obey  the  known  physical  laws.  In  fact, 
there  is  evidence  that  some  of  the  salts,  iron  for  example,  are  taken  up  as 
organic  compounds  (hematogens  of  Bunge).  The  activity  of  the  epithelial 
cells  is  to  be  taken  into  account,  even  in  the  absorption  of  salts. 

Water,  which  we  have  seen  is  not  absorbed  in  the  stomach,  is  readily 
absorbed  in  the  small  intestine.  Perhaps  the  bulk  of  water  taken  into 
the  system  is  absorbed  in  the  upper  part  of  the  small  intestine.  In  the  large 
intestine,  too,  it  is  absorbed  with  facility  and  in  considerable  quantities. 
The  content  of  the  bowel  is  still  quite  fluid  when  it  enters  the  ascending  colon, 
but  the  feces  are  quite  firm  on  discharge  from  the  rectum.  There  are  many 
analogies  by  which  we  may  suppose  a  controlling  influence  of  the  epithelium 
over  the  process  of  water  absorption.  Among  the  fishes  there  are  species, 
the  salmon  for  example,  in  which  the  blood  maintains  a  relatively  constant 
osmotic  pressure,  and  therefore  salt  content.  In  the  salmon  this  is  about 
the  same  as  that  of  human  blood.  The  blood  is  separated  in  the  gills  by 
an  extremely  thin  epithelium  from  the  water  in  which  the  animals  live,  yet 
these  fishes  go  with  impunity  from  sea- water,  with  two  and  a  half  times  more 
salt  than  the  blood,  to  fresh  water  with  practically  no  salt  at  all.  The  epi- 
thelium of  the  gills  permits  the  passage  of  oxygen,  but  it  does  not  permit 
the  diffusion  or  dialysis  of  the  salts  or  the  water  in  either  direction.  It  is 
possible  that  there  is  a  certain  amount  of  resistance  to  the  passage  of  water 
through  the  walls  of  the  stomach,  while  the  intestinal  epithelium  permits 
water  to  pass  readily. 

The  factors  active  in  absorption  are  under  searching  investigation  at  the 
present  time,  so  that  it  is  reasonable  to  hope  that  the  near  future  will  give 
a  more  exact  understanding  of  this  intricate  subject. 


420  ABSORPTION 

ABSORPTION  FROM  THE  SKIN,  THE  LUNGS,  ETC. 

The  dry  corneous  stratified  epithelium  covering  the  human  body  pos- 
sesses great  resistance  to  the  absorption  of  most  substances.  The  sebaceous 
secretion  keeps  the  surface  slightly  oily.  Watery  solutions  do  not  readily 
wet  the  surface  and  therefore  do  not  penetrate.  There  is  some  absorption 
of  water  on  prolonged  contact  with  the  skin,  but  the  amount  is  insignificant. 
Medicated  baths,  especially  hot  baths,  may  be  accompanied  by  some  slight 
absorption  of  the  substances  dissolved  in  the  waters,  though  it  must  be  con- 
fessed that  the  primary  good  effects  of  such  treatment  come  from  other 
sources. 

On  the  other  hand,  oily  substances  come  in  more  intimate  contact  with 
the  skin  and  penetrate  deeper  and  more  readily.  Therefore,  lotions  con- 
taining medicines  are  occasionally  applied  to  the  skin,  and  slow  but  gradual 
absorption  occurs.  The  volatile  oils  penetrate  the  skin  readily. 

The  epithelial  lining  of  the  lungs  seems  peculiarly  adapted  to  the  quick 
absorption  of  gases  and  volatile  substances.  This  is  illustrated  by  the 
rapidity  of  adjustment  of  the  body  to  variations  in  pressure  in  the  inert 
nitrogen  of  the  air  in  caisson  work  and  in  aviation.  The  volatile  anes- 
thetics, ether  and  chloroform,  penetrate  the  pulmonary  epithelium  with 
greatest  facility.  The  same  is  true  for  other  volatile  substances.  There  is 
increasing  evidence  that  most  substances  soluble  in  water  penetrate  the 
pulmonary  epithelium  with  facility,  and  that  this  may  in  the  future 
become  a  more  used  channel  for  administering  medicines. 

Solutions  injected  into  or  otherwise  brought  into  contact  with  the  sub- 
dermal  connective  tissue,  the  surface  of  the  body  of  a  muscle,  or  intra- 
muscularly, or  within  the  peritoneal  or  thoracic  cavities,  very  quickly  pass 
into  the  general  circulation.  They  are  practically  injected  into  the  lymph- 
atic intercellular  spaces  in  these  instances  and,  of  course,  are  very  readily 
carried  through  the  lymphatic  vessels,  figures  280  and  282,  to  the  thoracic 
duct  and  into  the  blood.  Readily  diffusible  substances,  however,  pass 
directly  through  the  blood  capillary  walls  and  do  not  necessarily  follow  the 
longer  lymph  channel  route.  Comparing  the  rapidity  of  absorption  in  the 
cases  mentioned,  that  from  the  muscle  is  most  rapid,  a  fact  of  medical  im- 
portance in  the  use  of  the  hypodermic  needle  for  the  giving  of  medicines 
in  emergency. 


CHAPTER  X. 
EXCRETION. 

EVERY  substance  taken  into  the  body,  in  whatever  form,  must,  in  the 
end,  be  cast  off  again,  no  matter  how  great  the  change  that  may  be  wrought 
during  its  sojourn.  We  have  already  found  that  in  the  lungs  the  expired 
air,  and  in  the  intestine  the  feces,  carry  from  the  body  waste  matters  of  no 
further  use.  We  have  now  to  find  that  the  kidneys,  separating  the  urine, 
and  the  skin,  separating  the  sweat  and  the  sebum,  are  likewise  channels  by 
which  the  body  throws  off  water,  salts,  and  broken-down  organic  matters 
of  no  further  use  to  the  organism.  Of  these  two  organs,  the  skin  and  the 
kidney,  the  latter  is  by  far  the  more  important  in  so  far  as  the  quantity  and 
complexity  of  its  secretion  is  concerned. 

STRUCTURE  AND  FUNCTION  OF  THE  KIDNEYS. 

General  Structure. — The  kidneys  are  two  in  number,  and  are  situated 
deeply  in  the  lumbar  region  of  the  abdomen  on  either  side  of  the  spinal  col- 
umn behind  the  peritoneum.  They  correspond  in  position  to  the  last  two 
dorsal  and  two  upper  lumbar  vertebrae,  the  right  slightly  below  the  left  in 
consequence  of  the  position  of  the  liver  on  the  right  side  of  the  abdomen. 
They  are  about  4  inches  long,  2\  inches  broad,  and  ij  inches  thick.  The 
weight  of  each  kidney  is  about  4}  ounces,  140  grams. 

On  dividing  a  kidney  into  two  equal  parts  by  a  section  carried  through 
its  long  convex  border,  figure  286,  the  main  part  of  its  substance  is  seen  to 
be  composed  of  two  chief  portions  called,  respectively,  cortical  and  medullary, 
the  latter  being  also  sometimes  called  pyramidal,  from  the  fact  of  its  being 
composed  of  about  a  dozen  conical  bundles  of  uriniferous  tubules,  each  bun- 
dle forming  what  is  called  a  pyramid.  The  upper  part  of  the  ureter,  or  duct 
of  the  organ,  is  dilated  into  the  pelvis;  and  this,  again,  after  separating  into 
two  or  three  principal  divisions,  is  finally  subdivided  into  8  to  12  smaller 
portions,  calyces,  each  of  which  receives  the  pointed  extremity  or  papilla  of 
a  pyramid.  Sometimes,  however,  more  than  one  papilla  is  received  by  a 
calyx. 

The  kidney  is  a  compound  tubular  gland.  Both  its  cortical  and  its 
medullary  portions  are  composed  essentially  of  numerous  tubes,  the  tubuU 
uriniferi,  which  begin  at  the  opening  on  the  Malpighian  pyramid  and,  after 
a  devious  course,  end  in  the  capsule  of  the  glomerulus. 

421 


422 


EXCRETION 


Tubuli  Uriniferi. — The  tubuli  uriniferi,  figure  287,  are  composed 
of  a  nearly  homogeneous  membrane,  and  are  lined  internally  by  epithelium. 
They  vary  considerably  in  size  in  different  parts  of  their  course,  but  are, 
on  an  average,  about  40^  in  diameter,  and  are  found  to  be  made  up  of  several 
distinct  sections.  The  first  section  or  part  to  be  identified  is  the  Mal- 
pighian,  or  Bowman's  capsule,  figure  287.  It  is  composed  of  a  hyaline  mem- 
brana  propria,  thickened  by  a  varying  amount  of  fibrous  tissue,  and  lined  by 

a  b 


FIG.  286. — Longitudinal  Section  of  Kidney  through  Hilum.  a,  Cortical  pyramid;  6, 
medullary  ray;  c,  medulla;  d,  cortex;  e,  renal  calyx;/,  hilum;  g,  ureter;  h,  renal  artery;  », 
obliquely  cut  tubules  of  medulla;  j  and  k,  renal  arches;  /,  column  of  Bertini;  m,  connective 
tissue  and  fat  surrounding  renal  vessels;  n,  medulla  cut  obliquely;  o,  papilla;  p,  medullary 
pyramid.  (Merkel-Henle.) 

flattened  nucleated  epithelial  plates.  This  capsule  is  the  dilated  extremity  of 
the  uriniferous  tubule  which  is  invaginated  to  receive  the  glomerulus  of  con- 
voluted capillary  blood  vessels.  The  invaginated  portion  of  the  tubule  is  of 
particular  importance  since  it  is  the  membrane  through  which  a  large  part 
of  the  urine  is  secreted.  The  glomerulus  is  connected  with  an  efferent  and 
an  afferent  blood-vessel.  The  Malpighian  capsule  is  connected  by  a  con- 
stricted neck,  figure  287,  TV,  with  the  proximal  convoluted  tubule.  This  forms 
several  distinct  curves  and  is  lined  with  short  columnar  cells.  The  tube 


TUBULI    URINIFERI 


423 


next  passes  almost  vertically  downward  toward  the  medulla,  forming  the 
spiral  tubule,  still  within  the  cortex  of  the  kidney,  which  is  of  much  the  same 
diameter.  The  loop  of  Henle,  L,  in  the  medulla,  is  a  very  narrow  tube  lined 
with  flattened  nucleated  cells.  Passing  vertically  upward  from  the  loop  of 


LABYRINTH     \MF.IXRAY\LABYR. 


PC  I  v  it 

FIG.  287. — Scheme  of  Uriniferous  Tubule  and  of  the  Blood  Vessels  of  the  Kidney, 
Showing  Their  Relation  to  Each  Other  and  to  the  Different  Parts  of  the  Kidney.  G, 
Glomerulus;  BC,  Bowman's  capsule;  N,  neck,  PC,  proximal  convoluted  tubule;  5,  spiral 
tubule;  D,  descending  arm  of  Henle's  loop;L,  Henle's  loop;  A,  ascending  arm  of  Henle's 
loop;  IDC,  distal  convoluted  tubule;  AC,  arched  tubule;  SC,  straight  collecting  tubule; 
ED,  duct  of  Bellini;  A,  arcuate  artery,  and  V,  arcuate  vein,  giving  off  interlobular  vessels 
to  cortex  and  vasa  recta  to  medulla;  a,  afferent  vessel  of  glomerulus;  e,  efferent  vessel  of 
glomerulus;  c,  capillary  network  in  cortical  labyrinth;  s,  stellate  veins;  vr,  vasa  recta  and 
capillary  network  of  medulla.  (Pearsol.) 

Henle,  the  tubule  varies  somewhat  in  histological  character,  but  the  irregular 
tubule  and  the  distal  convoluted  tube,  identical  in  all  respects  with  the  proxi- 
mal convoluted  tube,  are  to  be  noted.  The  proximal  convoluted  tubes 
pass  into  the  curved  and  straight  collecting  tubes,  the  latter  running 
vertically  downward  to  the  papillary  layer,  and,  joining  with  other  collecting 


424 


EXCRETION 


tubes,  form  larger  ducts  which  finally  open  at  the  apex  of  the  papilla.     These 
collecting  tubes  are  lined  with  nucleated  columnar  or  cubical  cells. 

Renal  Blood  Supply. — The  renal  artery  divides  into  several  branches 
which  pass  in  at  the  hilus  of  the  kidney  and  are  covered  by  a  fine  sheath 
of  areolar  tissue  derived  from  the  capsule.  They  enter  the  substance  of  the 
organ  chiefly  in  the  intervals  between  the  papillae  and  at  the  junction  between 
the  cortex  and  the  boundary  layer.  The  main  branches  then  pass  almost 
horizontally,  forming  more  or  less  complete  arches  and  giving  off  branches 


FIG.  288. — From  a  Vertical  Section  through  the  Kidney  of  a  Dog,  the  Capsule  of  which 
is  Supposed  to  be  on  the  Right,  a,  The  capillaries  of  the  Malpighian  capsule,  the  glomerulus, 
are  arranged  in  lobules;  n,  neck  of  capsule;  c,  convoluted  tubes  cut  in  various  directions;  b, 
irregular  tubule;  d,  e,  and  /are  straight  tubes  running  toward  capsules  forming  a  so-called 
medullary  ray;  d,  collecting  tube;  e,  spiral  tube;  /,  narrow  section  of  ascending  limb. 
X  380.  (Klein  and  Noble  Smith.) 

upward  to  the  cortex  and  downward  to  the  medulla.  The  former  are  for 
the  most  part  straight;  they  pass  almost  vertically  to  the  surface  of  the  kidney, 
giving  off  laterally  in  all  directions  longer  and  shorter  branches,  which  ulti- 
mately supply  the  glomerulus.  The  small  afferent  artery,  figures  287,  a, 
290,  d,  which  enters  the  Malpighian  capsule,  breaks  up  in  the  interior  into 
a  dense  convoluted  and  looped  capillary  plexus,  which  is  ultimately  gathered 
up  again  into  several  efferent  vessels,  comparable  to  minute  veins,  which 
leave  the  capsule  at  one  or  more  places  near  the  point  at  which  the  afferent 
artery  enters  it.  On  leaving,  they  do  not  immediately  join  other  small 
veins  as  might  have  been  expected,  but  again  break  up  into  a  second  set 


RENAL  BLOOD    SUPPLY 


425 


of  capillary  vessels  which  form  an  interlacing  network  around  the  urinif- 
erous  tubules.  This  second  capillary  plexus  terminates  in  small  veins 
which,  by  union  with  others,  help  to  form  the  radicles  of  the  renal  vein. 
These  form  venous  arches  corresponding  to  the  arterial  arches  situated 
between  the  medulla  and  cortex. 

Thus,  in  the  kidney,  the  blood  entering  by  the  renal  artery  traverses 
two  sets  of  capillaries  before  emerging  by  the  renal  vein,  an  arrangement 
which  may  be  compared  to  the  portal  system. 

The  tuft  of  vessels  within  the  Malpighian  capsule  in  the  course  of  de- 
velopment has  been  thrust  into  the  dilated  extremity  of  the  urinary  tubule, 
which  finally  completely  invests  it.  Thus  within  the  Malpighian  capsule 


FIG.  289. — Transverse  Section  of  a  Renal  Papilla,  a,  Large  tubes  or  papillary  ducts; 
b,  c,  and  d,  smaller  tubes  of  Henle;  e,  f,  blood  capillaries,  distinguished  by  their  flatter 
epithelium.  (Cadiat.) 

there  are  two  layers  of  squamous  epithelium,  a  parietal  layer  lining  the  cap- 
sule proper,  and  a  visceral  or  reflected  layer  immediately  covering  the  vas- 
cular tuft,  figure  290,  and  sometimes  dipping  down  into  its  interstices.  This 
reflected  layer  of  epithelium  is  readily  seen  in  young  subjects,  but  cannot 
always  be  demonstrated  in  the  adult,  figures  290  and  291. 

The  vessels  which  enter  the  medullary  layer  break  up  into  smaller  arte- 
rioles,  which  form  a  fine  arterial  meshwork  around  the  tubes  of  the  papillary 
layer  and  end  in  a  similar  plexus  from  which  the  venous  radicles  arise.  The 
vessels  do  not  form  a  double  set  of  capillaries. 

Besides  the  small  afferent  arteries  of  the  Malpighian  bodies  there  are, 
of  course,  others  which  are  distributed  in  the  ordinary  manner,  for  the  nutri- 
tion of  the  different  parts  of  the  organ;  and  there  are  numerous  straight 


426 


EXCRETION 


vessels,  the  vasa  recta,  in  the  pyramids  between  the  tubes.  Some  of  these 
are  branches  of  the  vasa  efferentia  from  Malpighian  bodies,  and  therefore 
comparable  to  the  venous  plexus  around  the  tubules  in  the  cortical  portion, 
while  others  arise  directly  as  small  branches  of  the  renal  arteries. 


FIG.  290. — Malpighian  Capsule  and  Tuft  of  Capillaries,  Injected  through  the  Renal 
Artery  with  Colored  Gelatin,  a,  Glomerular  vessels;  b,  capsule;  c,  anterior  capsule;  d, 
glomerular  artery;  e,  efferent  veins;/,  epithelium  of  tubes.  (Cadiat.) 


FIG.  291. — Diagrams  Illustrating  Stages  in  the  Development  of  the  Malpighian  Cap- 
sule. In  i  and  2  the  developing  blood  vessel  is  approaching  the  blind  end  of  the  capsule. 
In  3  the  tubule  is  beginning  to  invaginate  and  enclose  the  capillary.  In  4  and  5  later 
stages  are  shown.  The  cells  forming  the  two  layers  of  the  capsule  grow  very  thin.  (Bailey.) 

Renal  Nerves. — Vaso-constrictor  and  vaso-dilator  nerves  are  sup- 
plied to  the  blood  vessels  of  the  kidney,  but  no  clearly  denned  secretory 
nerves  have  yet  been  demonstrated  for  the  organ.  The  vascular  nerves 
arise  out  of  the  anterior  spinal  roots  (Bradford),  chiefly  the  eleventh  to  the 


THE    URINE  427 

thirteenth  dorsal  nerves.  They  reach  the  kidney  by  way  of  the  splanchnic 
nerves  and  the  renal  plexus  to  the  renal  artery  along  which  they  run  into 
the  substance  of  the  kidney.  Berkely  has  demonstrated  nerve  plexuses 
about  the  arterioles  and  around  Bowman's  capsule.  Terminal  knob-like 
endings  of  nerve  fibrils  were  shown.  Some  authors  have  claimed  renal  vaso- 
constriction  following  vagus  stimulation,  but  the  fact  seems  not  to  be  uni- 
versally admitted. 

The  Ureters  and  Urinary  Bladder.— The  duct  of  each  kidney,  the 
ureter,  is  a  tube  about  the  size  of  a  goose-quill  and  from  twelve  to  sixteen 
inches  in  length.  It  is  continuous  above  with  the  pelvis  of  the  kidney, 
and  ends  below  by  obliquely  perforating  the  walls  of  the  bladder  and 
opening  on  its  internal  surface.  It  has  three  principal  coats,  an  outer 
fibrous,  a  middle  muscular,  of  which  the  fibers  are  unstriped  and  arranged 
in  three  layers.  The  fibers  of  the  central  layer  are  circular,  and  those  of  the 
other  two  layers  longitudinal  in  direction.  It  has  an  internal  mucous  lining 
continuous  with  that  of  the  pelvis  of  the  kidney  above  and  the  lining  of  the 
urinary  bladder  below.  The  urinary  bladder,  which  forms  a  receptacle  for 
the  temporary  lodgment  of  the  urine  in  the  intervals  of  its  expulsion  from 
the  body  is  more  or  less  pyriform.  Its  widest  part,  which  is  situated 
above  and  behind,  is  termed  thefundus;  and  the  narrow  constricted  portion 
in  front  and  below,  by  which  it  becomes  continuous  with  the  urethra,  is 
called  its  cervix  or  neck.  It  is  constructed  of  four  principal  coats:  serous, 
muscular,  areolar  or  submucous,  and  mucous.  The  fibers  of  the  muscular 
coat  deserve  special  mention.  They  are  unstriped,  are  arranged  in  three 
principal  layers,  of  which  the  external  and  internal  have  a  general  longitu- 
dinal, and  the  middle  layer  a  circular,  direction.  The  latter  are  especially 
developed  around  the  cervix  of  the  organ,  and  are  described  as  forming  a 
sphincter  vesicae.  The  mucous  membrane  is  provided  with  mucous  glands, 
which  are  more  numerous  near  the  neck  of  the  bladder. 

The  bladder  is  well  provided  with  blood  and  lymph  vessels  and  with 
nerves.  The  latter  are  from  the  sacral  plexus  (spinal)  and  hypogastric 
plexus  (sympathetic).  Ganglion  cells  are  found,  here  and  there,  in  the 
course  of  the  nerve  fibers. 

THE  URINE. 

Quantity  and  General  Properties. — Healthy  urine  is  a  perfectly 
transparent  amber-colored  liquid,  with  a  peculiar  but  not  disagreeable  odor, 
a  bitterish  salty  taste,  and  a  specific  gravity  of  from  i  .020  to  i  .025.  The 
urine  consists  of  water  holding  in  solution  certain  organic  and  saline  matters 
as  its  ordinary  constituents,  and  occasionally  various  other  matters.  Some 
of  the  latter  are  indications  of  diseased  states  of  the  system  and  others  are 
derived  from  unusual  articles  of  food  or  drugs  taken  into  the  stomach. 


428  EXCRETION 

The  total  quantity  of  urine  passed  in  twenty-four  hours  is  influenced 
by  numerous  circumstances.  In  adults  of  average  size  and  medium  ac- 
tivity the  daily  amount  of  urine  may  be  given  as  from  1,200  cc.  to  1,500  cc. 
In  Chittenden's  recent  observations  on  nine  athletic  students  and  on  eight 
soldiers  the  average  daily  output  of  urine  through  a  period  of  about  five 
months  was  for  the  students  1,215  cc.  with  average  specific  gravity  of  i  .020, 
and  for  the  soldiers  1,042  cc.  with  specific  gravity  of  i  .023. 

GENERAL  CHEMICAL  COMPOSITION  OF  THE  URINE. 

Water 967 

Solids: 

Urea 14.230 

Other  nitrogenous  crystalline  bodies: 

Uric  acid,  principally  in  the  form  of  alkaline  urates, 

a  trace  only  free 

Kreatinin,  xanthin,  hypoxanthin f 

Hippuric  acid 

Mucus,  pigments,  and  ferments 

Salts: 

Inorganic: 

Principally  sulphates,  phosphates,  and  chlorides 
of  sodium  and  potassium,  with  phosphates 
of  magnesium  and  calcium,  traces  of  silicates 
Organic: 

Lactates,  oxalates,  acetates,  butyrates  and  for- 
mates, which  appear  only  occasionally.  .  .  .  j  -  33 

Sugar a  trace  sometimes. 

Gases  (nitrogen  and  carbon  dioxide  principally). 


1,000 


Reaction.— The  normal  reaction  of  the  urine  is  slightly  acid.  This 
acidity  is  due  to  carbon  dioxide  and  to  acid  phosphate  of  sodium,  and  is  less 
marked  soon  after  meals.  After  a  time,  varying  in  length  according  to  the 
temperature,  the  reaction  becomes  strongly  alkaline  from  the  change  of 
urea  into  ammonium  carbonate,  due  to  the  presence  of  one  or  more  specific 
micro-organisms  (micrococcus  urea).  In  the  process  of  fermentation  the 
urea  takes  up  two  molecules  of  water,  a  strong  ammoniacal  and  fetid  odor 
appears,  and  there  are  deposits  of  triple  phosphates  and  alkaline  urates. 
This  does  not  occur  unless  the  urine  is  freely  exposed  to  the  air,  or,  at  least, 
until  air  has  had  access  to  it. 

In  most  herbivorous  animals  the  urine  is  alkaline  and  turbid.  The 
difference  depends  not  on  any  peculiarity  in  the  mode  of  secretion,  but  on 
the  difference  in  the  food  on  which  the  two  classes  of  animals  subsist.  For 
when  carnivorous  animals,  such  as  dogs,  are  restricted  to  a  vegetable  diet, 
their  urine  becomes  pale,  turbid,  and  alkaline  like  that  of  herbivorous 
animals,  while  the  urine  voided  by  the  herbivora,  e.g.,  rabbits,  fed  for  some 


VARIATIONS    IN    THE    CONSTITUENTS    OF    URINE  429 

time  exclusively  upon  animal  substances,  presents  the  acid  reaction  and 
other  qualities  of  the  urine  of  carnivora,  and  its  ordinary  alkalinity  is  again 
restored  only  on  the  substitution  of  a  vegetable  for  the  animal  diet.  Human 
urine  is  not  usually  rendered  alkaline  by  vegetable  diet,  but  it  becomes  so 
after  the  free  use  of  alkaline  medicines  or  of  the  alkaline  salts  with  carbonic 
or  vegetable  acids.  These  latter  are  changed  into  alkaline  carbonates 
previous  to  elimination  by  the  kidneys. 

Specific  Gravity  of  Urine. — The  average  specific  gravity  of  the  human 
urine  is  about  i  .020  to  i  .025.  The  relative  quantity  of  water  and  of  solid 
constituents  of  which  it  is  composed  is  materially  influenced  by  the  condition 
and  occupation  of  the  body  during  the  time  at  which  it  is  secreted;  by  the 
length  of  time  which  has  elapsed  since  the  last  meal;  by  the  amount  of  water 
taken;  and  by  several  other  less  important  circumstances.  The  morning 
urine  is  the  best  adapted  for  analysis  in  health,  since  it  represents  the  simple 
secretion  unmixed  with  the  elements  of  food  or  drink.  If  it  is  not  used  the 
whole  of  the  urine  passed  during  the  period  of  twenty-four  hours  should  be 
taken.  The  specific  gravity  of  the  urine  may  thus,  consistently  with  health, 
range  widely  on  both  sides  of  the  usual  average.  It  may  vary  from  i  .015 
in  the  winter  to  1.025  in  the  summer;  but  variations  of  diet  and  exercise, 
and  many  other  circumstances,  may  make  even  greater  differences  than 
these.  The  variations  may  be  extreme  in  disease,  sometimes  decreasing  in 
chronic  nephritis  to  i .  004,  and  frequently  increasing  in  diabetes,  when  the 
urine  is  loaded  with  sugar,  to  i .  050  or  even  to  i .  060. 

AVERAGE  DAILY  QUANTITY  OF  THE  CHIEF  URINARY  CONSTITUENTS     (MODI- 
FIED FROM  PARKES.) 

Per  kilo  of 
Body  weight. 

Water 1,500.          c.c.  23.0000  grams 

Solids 72 .           grams  o .  8800  grams 

Urea 33  . 180   grams  o  .  5000  grams 

Kreatinin .910   grams  0.0140  grams 

Uric  acid .  555   grams  o  .0084  grams 

Hippuric  acid .400   grams  o  .0060  grams 

Pigment  and  extractives 10.000    grams  o.  1510  grams 

Sulphuric  acid 2.012    grams  0.0480  grams 

Phosphoric  acid 3-164    grams  0.0305  grams 

Chlorine 7  .000   grams  o .  1260  grams 

Ammonia -77°   grams 

Potassium 2  .  500   grams 

Sodium ii.  090   grams 

Calcium .260   grams 

Magnesium .  207    grams 

Variations  in  the  Constituents  of  Urine. — Most  of  the  constituents 
are,  even  in  health,  liable  to  variations  from  the  proportions  given  in  the 
above  table.  The  variations  of  the  quantity  of  water  in  different  seasons 


430  EXCRETION 

and  according  to  the  quantity  of  drink  and  exercise  have  just  been  men- 
tioned. The  water  of  the  urine  is  also  liable  to  be  influenced  by  the  condi- 
tion of  the  nervous  system,  being  sometimes  greatly  increased,  e.g.,  in  hysteria 
and  in  some  other  nervous  affections,  and  at  other  times  diminished.  The 
increase  in  water  may  be  either  attended  with  an  augmented  quantity  of 
solid  matter  in  some  diseases,  as  in  ordinary  diabetes,  or  may  be  nearly  the 
sole  change,  as  in  the  affection  termed  diabetes  insipidus.  A  febrile  con- 
dition almost  always  diminishes  the  quantity  of  water;  and  a  like  diminution 

is  caused  by  any  affection  which  draws  off 
a  large  quantity  of  fluid  from  the  body 
through  any  other  channel  than  that  of  the 
kidneys,  e.g.,  the  bowels  or  the  skin. 

In  disease  or  after  the  ingestion  of  special 
foods,  various  abnormal  substances  occur  in 
urine,  of  which  the  following  may  be  men- 
tioned:     Seralbumin,       serum       globulin, 
enzymes    (apparently   some  are  present  in 
health  also),   proteoses,   blood,   sugar,   bile 
acids  and  pigments,  casts,  fats,  various  salts 
FIG.  292. — Crystals  of  Urea.        taken  as  foods  or  as  medicines,  micro-organ- 
isms of  various  kinds. 

The  Nitrogenous  Substances  in  Urine. — The  nitrogenous  waste  prod- 
ucts which  are  formed  in  the  body  in  the  metabolism  of  the  protein  foods 
are  ultimately  eliminated  chiefly  through  the  kidney,  to  some  extent  through 
the  bowel,  and  slightly  through  the  skin.  The  total  nitrogen  in  the  urine 
and  in  the  feces  multiplied  by  the  factor  6.25  is  a  measure  of  the  nitrogenous 
foods,  i.e.,  proteins,  metabolized  by  the  body.  The  nitrogen  excreted  in  the 
urine  is  in  the  form  of  urea  87  . 5  per  cent.,  ammonia  4.3  per  cent.,  kreatinin 
3 . 6  per  cent.,  uric  acid  o .  8  per  cent.,  and  undetermined  forms  3.73  per  cent., 
according  to  Folin.  The  total  quantity  of  nitrogen  eliminated  in  all  these 
forms  per  day  is  given  as  about  18  grams.  In  Chittenden's  experiments 
this  quantity  is  reduced  to  as  low  as  6  grams  or  even  less  per  day. 

Urea. — Urea,  CON2H4,  is  the  principal  solid  constituent  of  the  urine, 
forming  nearly  one-half  of  the  total  quantity.  It  is  also  the  most  important 
ingredient,  since  it  is  the  chief  form  in  which  the  waste  nitrogen  which  is 
derived  from  protein  metabolism  is  excreted  from  the  body. 

Properties. — Urea,  like  other  solid  constituents  of  the  urine,  exists  in  a 
state  of  solution.  When  in  the  solid  state,  it  appears  in  the  form  of  delicate 
silvery  acicular  crystals,  which,  under  the  microscope,  are  seen  as  four- 
sided  prisms,  figure  292.  It  readily  combines  with  some  acids,  like  a  weak 
base,  and  may  thus  be  conveniently  procured  in  the  form  of  crystals  of  nitrate 
or  oxalate  of  urea,  figures  293  and  294. 

Urea  is  colorless  when  pure;  when  impure  it  may  be  yellow  or  brown. 


THE    FORMATION    OF   UREA 


431 


It  is  without  odor  and  of  a  cooling  niter-like  taste.  It  has  neither  an  acid 
nor  an  alkaline  reaction,  and  deliquesces  in  a  moist  and  warm  atmosphere. 
At  15°  C.  it  requires  for  its  solution  less  than  its  own  weight  of  water.  It  is 
soluble  in  all  proportions  of  boiling  water,  and  requires  five  times  its  weight 
of  cold  alcohol  for  its  solution.  It  is  insoluble  in  ether.  At  120°  C.  it  melts, 
and  at  a  still  higher  temperature  decomposes. 

Urea  is  decomposed  by  sodium  hypochlorite  or  hypobromite  or  by  nitrous 
acid,  with  evolution  of  nitrogen.  It  forms  compounds  with  acids,  of  which 
the  chief  are  urea  hydrochloride,  CON2H4.HC1;  urea  nitrate,  CON^H4.- 
HNO3;  and  urea  phosphate,  CON2H4.H3PO4.  It  forms  compounds  with 


FIG.  293.— Crystals  of  Urea  Nitrate.  FIG.  294.— Crystals  of  Urea  Oxalate. 

metals  such  as  HgO .  CON2H4,  (or  with  silver,  CON2H2Ag2).  Urea  is  iso- 
meric  with  ammonium  cyanate,  NH4CNO,  and  was  first  prepared  artificially 
from  that  substance. 

The  Formation  of  Urea. — Proteins  in  the  body  have  their  nitrog- 
enous moiety  split  off  as  ammonia,  by  what  Folin  considers  essentially 
a  series  of  hydrolytic  cleavages;  this  is  then  built  up  into  urea,  as  described 
more  fully  in  the  chapter  on  Metabolism.  This  last  step  is  essentially  a 
synthetic  process  which,  from  the  fact  that  ammonium  carbonate  introduced 
into  the  blood  is  eliminated  as  urea,  may  be  supposed  to  occur  as  follows: 

/O.NH4  /NH2  /HN2 

0:C<  — H20     ->  0:C<  -H2O  ->  O:C< 

XO.NH4  X).NH4  XNH2 

Ammonium  Ammonium  Urea 

Carbonate  Carbamate 

Urea  is  present  in  varying  amounts  in  all  organs  and  fluids  of  the  body,  as 
shown  by  the  following  determinations  of  Schoendorff  on  the  dog: 


Organ 

Blood 0.116 

Muscle o .  080 

Kidney o .  670 

Liver 0.112 

Heart , o.  173 

Brain o.  128 

Spleen 0.122 


432 


EXCRETION 


It  has  oeen  proven  tftat  the  kidney  does  not  form  urea;  in  fact  the  kid- 
neys may  be  removed  from  the  body,  and  urea  will  continue  to  accumulate 
in  the  blood.  Urea  is  formed  chiefly  in  the  liver,  but  may  in  part  be  con- 
structed in  other  organs,  as  described  more  fully  on  page  408.  It  follows 
that  the  kidney  is  only  the  channel  for  the  elimination  of  this  nitrogenous 
compound. 

Decomposition  of  the  urea  with  development  of  ammonium  carbonate 
takes  place  from  the  action  of  bacteria  (micrococcus  ureae)  when  urine  is 
kept  for  some  days  after  being  voided,  which  explains  the  ammoniacal  odor 
then  evolved.  The  urea  is  sometimes  decomposed  before  it  leaves  the  blad- 
der, when  the  mucous  membrane  is  diseased  and  the  mucus  secreted  by  it  is 
abundant;  but  decomposition  does  not  occur  unless  atmospheric  germs  have 
had  access  to  the  urine. 

Quantity  Excreted. — The  quantity  of  urea  excreted  is,  like  that  of  the  urine 
itself,  subject  to  considerable  variation.  For  a  healthy  adult  about  30  grams 
per  day  may  be  taken  as  rather  a  high  average.  Its  percentage  in  healthy 
urine  is  from  2  to  2.5.  Its  amount  is  materially  influenced  by  diet,  being 
greater  on  a  diet  of  high  protein  content.  The  quantity  of  urea  excreted 
by  children,  relatively  to  their  body  weight,  is  much  greater  than  by  adults; 
thus  the  quantity  of  urea  excreted  per  kilogram  of  weight  was  found  to  be, 
in  a  child,  o .  8  gram;  in  an  adult  only  o .  4  gram.  Regarded  in  this  way,  too, 
the  excretion  of  carbonic  acid  gives  similar  results,  the  proportions  in  the 
child  and  adult  being  as  82  to  34. 

Uric  Acid. — Uric  acid,  C5H4N4O3,  see  page  410,  is  present  in  the  urine 
of  man  and  other  animals.  In  birds  and  reptiles  uric  acid  or  its  salts  is  the 
chief  form  in  which  nitrogen  is  eliminated  from  the  body. 

Properties. — Uric  acid  is  a  colorless,  crystalline  compound  of  the  purin 
group,  figure  295.  It  is  odorless  and  tasteless.  It  is  very  slightly  soluble  in 
water,  quite  insoluble  in  alcohol  and  ether,  and  freely  soluble  in  solutions 
of  the  alkaline  carbonates  and  other  salts. 

A  study  of  the  elimination  of  nitrogen  in  birds,  i.e.,  geese,  has  shown  that 
uric  acid,  like  urea  in  mammals,  is  formed  largely  in  the  liver  from  antecedent 
protein  nitrogen.  In  man  the  elimination  of  uric  acid  (o .  3  to  o .  6  gram  per 
diem)  is  more  or  less  constant  and  characteristic  for  the  individual;  it  in- 
creases or  decreases  somewhat  with  the  nucleoprotein  and  purin  content 
of  the  daily  diet.  This  observation  has  led  to  the  inference  that  uric-acid 
nitrogen  is  derived  from  nuclear  metabolism,  page  94. 

A  certain  amount  of  nuclear  purin  is  taken  in  the  food,  called  exogenous 
purin  to  distinguish  it  from  the  endogenous  purin  that  is  derived  from  the 
body  nuclear  metabolism. 

Other  representatives  of  the  purin  group  are  adenin,  guanin,  xanthin, 
hypoxanthin,  etc.  Chemically,  caffeine  from  coffee  is  a  trimethyl  xanthin. 

The  most  common  form  in  which  uric  acid  is  deposited  in  urine  is  that 


PIGMENTS 


433 


of  a  brownish  or  yellowish  powdery  substance,  consisting  of  amorphus 
ammonium  or  sodium  urate.  Urate  sediments  are  commonly  deposited  on 
cooling  the  urine;  they  are  redissolved  on  warming  it  slightly.  When  de- 
posited in  crystals,  uric  acid  is  most  frequently  obtained  in  rhombic  or  dia- 
mond-shaped laminae,  but  other  forms  are  not  uncommon,  figure  295. 
When  deposited  from  urine,  the  crystals  are  generally  more  or  less  deeply 
colored,  from  being  combined  with  the  coloring  principles  of  the  urine. 

Hippuric  Acid.— This  compound,  C6H5.CO.NH.CH2COOH,  has  long 
been  known  to  exist  in  the  urine  of  herbivorous  animals  in  combination  with 
sodium  It  also  exists  naturally  in  the  urine  of  man,  in  a  quantity  equal  to, 


FIG.  295. — Various  Forms  of  Uric  Acid  Crystals.        FIG.  296. — Crystals  of  Hippuric  Acid. 

or  rather  exceeding,  that  of  the  uric  acid.  The  quantity  excreted  is  increased 
by  a  vegetable  diet. 

Hippuric  acid  appears  to  be  formed  in  the  body  from  benzoic  acid.  The 
benzoic  acid  unites  with  glycocoll,  and  hippuric  acid  and  water  are  formed 
thus: 

C6H5.COOH  +  CH2.NH2.COOH  =  C6H5.CO.NH.CH2.COOH  +  H2O. 

Benzoic  acid  Glycin  Hippuric  acid 

Hippuric  acid  is  the  first  substance  which  was  demonstrated  to  be  synthet- 
ized  in  the  body. 

Kreatinin. — This  substance,  which  is  the  anhydride  of  kreatin,  is  pres- 
ent in  urine  in  a  remarkably  constant  quantity,  as  shown  recently  by  Folin's 
analyses.  Its  daily  excretion  quantity  is  from  i  to  1.5  grams  according  to 
the  amount  of  active  tissue  in  the  individual.  It  is  of  especial  importance 
as  a  measure  of  the  metabolism  of  muscle  protoplasm. 

Ammonia. — A  considerable  daily  quantity  of  ammonia  (about  o .  7  gram) 
in  combination,  as  chloride,  phosphate,  or  sulphate,  is  found  in  the  normal 
urine,  showing  that  this  is  an  important  method  of  nitrogen  elimination. 

Pigments. — The  pigments  of  the  urine  are  the  following:  i.  Urochrome. 
a  yellow  coloring  matter,  giving  no  absorption  band;  of  which  but  little  is 
known.  Urine  owes  its  yellow  color  mainly  to  the  presence  of  this  body. 
2.  Urobilin,  an  orange  pigment,  of  which  traces  may  be  found  in  nearly  all 


434 


EXCRETION 


urines,  and  which  is  especially  abundant  in  the  urines  passed  by  febrile 
patients.  It  is  characterized  by  a  well-marked  spectroscopic  absorption 
band  at  the  junction  of  green  and  blue.  Those  who  believe  urobilin  to 
be  identical  with  hydrobilirubin  suppose  that  the  bilirubin  is  reduced  by 
the  putrefactive  processes  in  the  intestines,  and  is  conveyed  in  its  reduced 
form  by  the  blood  stream  to  the  kidneys.  3.  Uroerythrin,  occasionally 
found. 

Mucus. — Mucus  sediment  in  the  urine  consists  principally  of  mucin  and 
of  the  epithelial  debris  from  the  mucous  surface  of  the  urinary  passages. 
Particles  of  epithelium,  in  greater  or  less  abundance,  may  be  detected  in  most 
samples  of  urine,  figure  297.  As  urine  cools,  the  mucus  is  sometimes  seen 
suspended  in  it  as  a  delicate  opaque  cloud,  but  generally  it  falls.  In  inflam- 
matory affections  of  the  urinary  passages,  especially  of  the  bladder,  mucus 
is  secreted  in  large  quantities. 


FIG.  287.  FIG.  298. 

FIG.  297. — Urinary  Deposit  of  Mucus,  etc. 

FIG.  298. — Urinary  Sediment  of  Triple  Phosphates  (large  prismatic  crystals)  and 
Urate  of  Ammonium,  from  urine  which  had  undergone  alkaline  fermentation. 

Saline  Matter.— Sulphuric  acid,  in  the  form  of  salts,  is  taken  in  very 
small  quantity  with  food.  Sulphur  is  also  a  constituent  part  of  the  protein 
molecule ;  hence  its  elimination,  like  that  of  nitrogen,  gives  a  certain  measure 
of  protein  metabolism.  It  is  excreted  as  inorganic  sulphates  of  sodium  and 
potassium,  and  as  eth  eal  sulphates,  compounds  of  phenol,  cresol,  skatol, 
i.e.,  cresol  sulphuric  acid  (C7H7OSO2OH),  etc. 

The  phosphoric  acid  in  the  urine  is  combined  partly  with  the  alkalies, 
partly  with  the  alkaline  earths, — about  four  or  five  times  as  much  with  the 
former  as  with  the  latter.  In  blood,  saliva,  and  other  alkaline  fluids  of  the 
body  phosphates  exist  in  the  form  of  alkaline,  neutral,  or  acid  salts.  In  the 
urine  they  are  acid  salts,  viz.,  the  sodium,  ammonium,  calcium,  and  magne- 
sium phosphates,  the  excess  of  acid  being  (Liebig)  due  to  the  appropriation 
of  the  alkali  with  which  the  phosphoric  acid  in  the  blood  is  combined,  by 
the  several  new  acids  which  are  formed  or  discharged  at  the  kidneys, 
namely  the  uric,  hippuric,  and  sulphuric  acids. 


OCCASIONAL    CONSTITUENTS    OF   URINE 


435 


The  phosphates  are  taken  largely  in  both  vegetable  and  animal  food. 
Some  are  excreted  at  once;  others  only  after  being  transformed  and  incor- 
porated with  the  tissues.  Calcium  and  magnesium  phosphates  form  the  prin- 
cipal earthy  constituents  of  bone,  and  from  the  decomposition  of  the  osseous 
tissue  the  urine  derives  a  quantity  of  this  salt.  The  decomposition  of  other 
tissues  also  furnishes  large  supplies  of  phosphorus  to  the  urine,  which  phos- 
phorus is  supposed,  like  the  sulphur,  to  be  united  with  oxygen,  and  then 
combined  with  bases.  The  quantity  is,  however,  liable  to  considerable 
variation.  The  earthy  phosphates  are  more  abundant  after  meals,  whether 
of  animal  or  vegetable  food,  and  are  diminished  after  long  fasting.  The 
alkaline  phosphates  are  increased  after  animal  food,  diminished  after  vege- 


FIG.  299. — Crystals  of  Cystin. 


FIG.  300. — Crystals  of  Calcium  Oxalate. 


table  food.  Phosphorus  uncombined  with  oxygen  appears,  like  sulphur,  to 
be  excreted  in  the  urine.  When  the  urine  undergoes  alkaline  fermentation, 
phosphates  are  deposited  in  the  form  of  a  urinary  sediment,  consisting  chiefly 
of  ammonio- magnesium  phosphate  (triple  phosphate),  figure  298. 

The  chlorine  of  the  urine  occurs  chiefly  in  combination  with  sodium. 
Next  to  urea,  sodium  chloride  is  the  most  abundant  solid  constituent  of  the 
urine.  As  the  chlorides  exist  largely  in  food,  and  in  most  of  the  animal 
fluids,  their  occurrence  in  the  urine  is  easily  understood. 

Occasional  Constituents  of  Urine. — Cystin,  C3H7NSO2,  figure  299, 
is  an  occasional  constituent  of  urine. 

Another  constituent  of  the  urine  is  oxalic  acid  (about  0.02  gram  per 
diem),  which  is  frequently  deposited  in  combination  with  calcium,  figure 
300,  as  a  urinary  sediment.  Pathologically,  oxalic  acid  is  found  to  be  in- 
creased in  diabetes  mellitus,  in  organic  diseases  of  the  liver,  and  in  various 
other  conditions  accompanied  by  derangement  of  the  oxidation  mechanism. 

Dextrose  and  albumin  are  sometimes  present  in  pathological  urine,  and  are 
of  particular  interest  from  the  clinical  point  of  view.  See  the  subject 
Internal  Secretions  of  the  Pancreas,  page  491. 


436  EXCRETION 

THE  METHOD  OF  EXCRETION  OF  URINE. 

The  secretion  of  urine  is  an  act  the  complexity  of  which  can  be  profitably 
discussed  only  after  a  clear  understanding  of  three  main  factors  which  have 
already  been  presented,  viz.,  the  chemical  composition  of  the  urine  secreted, 
the  structure  of  the  kidney  tubule  as  a  secreting  organ,  and,  finally,  the  chem- 
ical composition  of  the  blood  which  supplies  the  materials  to  the  kidney  for 
the  formation  of  the  urine.  The  substances  found  in  the  urine  are,  for  the 
most  part,  also  to  be  found  in  the  blood  plasma.  But  the  relative  percent- 
age composition  is  very  different.  The  amount  of  urea  in  the  blood  is  only  a 
fractional  part  as  concentrated  as  in  the  urine,  while  albumins  and  sugars, 
which  are  so  plentiful  in  the  blood,  are  normally  present  in  the  urine  only 
in  traces.  The  presence  of  the  glomerulus  with  its  special  vascular  supply, 
and  the  different  loops  of  the  tubule,  with  its  gland-like  epithelial  wall, 
would,  a  priori,  lead  one  to  suspect  special  functions  for  each. 

Theories  of  the  Secretion  of  Urine. — Bowman  in  1842,  wholly  on 
structural  grounds,  advanced  a  theory  of  urinary  secretion.  This  theory  was 
given  an  experimental  basis  and  elaborated  by  Heidenhain  and  generally 
bears  his  name. 

Heidenhain' 's  theory  is  stated  as  follows: 

i.  The  secretion  in  the  kidney  depends  upon  the  physiological  activity  of 
special  secreting  cells  which  are  of  two  kinds.  2.  The  first  type  of  cell  is 
represented  by  the  single  layer  of  epithelium  covering  the  glomerular  capil- 
laries. These  cells  secrete  especially  water  and  salts.  3.  The  second  type  of 
cell  is  represented  by  the  gland-like  epithelial  cells  which  form  the  convo- 
luted tubules  and  the  loop  of  Henle.  These  cells  secrete  the  urea,  uric  acid, 
and  other  specific  constituents  of  the  urine.  4.  The  activity  of  each  kind  of 
cell  is  influenced  by  the  chemical  composition  of  the  blood  and  by  the  flow 
of  blood  through  the  kidney.  5.  The  relative  secretory  activity  of  the  glomer- 
ular cells  and  the  tubule  cells  is  sufficient  to  account  for  the  variation  in  the 
chemical  composition  of  the  urine. 

Ludwig's  theory,  advanced  in  1844,  is  a  strictly  mechanical  theory  of 
urinary  secretion  based  on  experiments  which  he  presented,  i .  He  considered 
the  glomerulus  and  Bowman's  capsule  as  a  filtering  apparatus  in  which 
substances  present  in  the  blood  are  driven  through  the  epithelium  of  the 
capsule  into  the  renal  tubule  by  the  positive  pressure  of  the  blood  in  the 
glomerular  capillaries.  2.  This  very  dilute  urine  in  the  capsule  is  supposed 
to  be  concentrated  by  the  resorption  of  water  as  it  flows  down  the  tubule. 
Ludwig  originally  considered  this  resorption  of  water  an  imbibition  process 
in  which  the  greater  saturation  of  salts  in  the  blood  caused  water  to  be  taken 
up  through  the  renal  tubule  walls,  an  osmotic  process.  At  present  most 
observers  who  accept  the  view  that  filtration  takes  place  at  the  glomerulus 
explain  the  resorption  of  water  down  the  tubules  as  an  act  of  cellular 
resorption  or  secretion. 
27 


EXPERIMENTAL   OBSERVATIONS  437 

Experimental  Observations. — There  are  numerous  nerves  to  the 
kidney,  but  no  proven  secretory  influence  has  been  shown.  The  variations 
in  the  secretion  of  urine  that  follow  nervous  stimulation  are  quite  satisfactorily 
explained  by  the  changes  in  the  blood  flow. 

The  kidney  can  be  placed  in  an  onkometer  and  its  variation  in  volume 
measured  directly,  figures  301  and  302.  This  volume  measurement,  when 
taken  with  the  arterial  pressure,  gives  a  very  good  index  of  the  volume  of 
blood  flowing  through  the  kidney.  Now  when  the  kidney  is  inserted  in  an 
onkometer  and  the  urine  collected  from  the  ureter,  it  is  found  in  general  that 


FIG.  301. — Diagram  of  Roy's  Onkometer.  a,  Represents  the  kidney  enclosed  in  a 
metal  box,  which  opens  by  hinge/;  b,  the  renal  vessels  and  ducts.  Surrounding  the  kidney 
are  two  chambers  formed  by  membranes,  the  edges  of  which  are  firmly  fixed  by  being 
clamped  between  the  outside  and  inside  metal  capsules  (the  latter  not  represented  in  the 
figure),  the  two  being  firmly  screwed  together  by  screws  at  h,  and  on  the  opposite  side. 
The  membranous  chamber  below  is  filled  with  a  varying  amount  of  warm  oil,  according 
to  the  size  of  the  kidney  experimented  with,  through  the  opening  then  closed  with  the  plug  i. 
After  the  kidney  has  been  enclosed  in  the  capsule,  the  membranous  chamber  above  is  filled 
with  warm  oil  through  the  tube  e,  which  is  then  closed  by  a  tap  (not  represented  in  the 
diagram);  the  tube  d  communicates  with  a  recording  apparatus,  and  any  alteration  in  the 
volume  of  the  kidney  is  communicated  by  the  oil  in  the  tube  to  the  chamber  d  of  the  Onko- 
graph,  figure  302. 

the  greater  the  pressure  and  flow  of  blood  the  greater  the  secretion  of  urine, 
as  would  follow  if  the  glomerulus  were  a  filtering  mechanism.  However,  if 
the  renal  vein  is  partially  obstructed,  even  though  the  blood  pressure  be  in- 
creased, the  amount  of  urine  secreted  is  sharply  decreased.  If  the  vein  is 
completely  occluded,  the  secretion  of  urine  not  only  ceases  for  the  time  but 
does  not  immediately  begin  again  when  the  blood  pressure  and  flow  are  re- 
established. The  closure  of  the  vein  for  only  one  or  two  minutes  is  said  to 
stop  the  flow  of  urine  for  as  much  as  forty-five  minutes.  This  short  inter- 
ruption of  the  circulation  is  sufficient  to  bring  about  other  changes  in  the 


438  EXCRETION 

glomerular  epithelium,  for  it  now  excretes  albumin,  which  it  did  not  previ- 
ously let  pass.  Therefore,  it  is  not  pressure  merely  that  favors  the  secretion, 
there  must  also  be  an  efficient  flow  of  blood.  The  secretion  is  influenced 
especially  by  the  amount  of  blood  flowing  through  the  kidney  in  a  given  time. 
In  the  frog  the  kidney  has  a  double  blood  supply.  The  renal  artery 
supplies  the  glomeruli,  while  a  branch  of  the  renal-portal  vein  supplies  the 
tubules.  Nussbaum  ligated  the  renal  artery  in  one  kidney  of  the  frog,  while 
leaving  the  circulation  of  the  other  kidney  undisturbed.  He  found  that  the 
operated  kidney  secreted  little  or  no  urine,  but  that  it  could  be  made  to  secrete 
by  injections  of  urea,  but  not  by  injections  of  albumin  or  sugar  as  in  the  nor- 


FIG.  302. — Roy's  Onkograph,  or  Apparatus  for  Recording  Alterations  in  the  Volume 
of  the  Kidney,  etc.,  as  shown  by  the  onkometer.  a,  Upright,  supporting  recording  lever  /, 
which  is  raised  or  lowered  by  the  needle  b,  which  works  through/,  and  which  is  attached 
to  the  piston  e,  working  in  the  chamber  d,  with  which  the  tube  from  the  onkometer  com- 
municates. The  oil  is  prevented  from  being  squeezed  out  as  the  piston  descends,  by  a 
membrane,  which  is  clamped  between  the  ring-shaped  surfaces  of  the  cylinder  by  the  screw 
i  working  upward;  the  tube  h  is  for  filling  the  instrument. 

mal  kidney.  Ligation  of  the  renal-portal  vein,  which  supplies  the  tubules 
in  the  frog,  caused  a  decrease  in  the  quantity  of  the  secretion,  whereas,  ac- 
cording to  Ludwig's  view,  it  ought  to  have  increased  the  quantity,  since 
obviously  resorption  could  not  take  place  with  any  degree  of  efficiency.  In 
the  main,  the  evidence  is  in  favor  of  the  view  that  even  the  glomerular  epi- 
thelium does  not  filter  merely,  but  that  it,  as  living  protoplasm,  regulates 
and  controls  the  quantity  and  kind  of  material  passing  through  it. 

Microchemical  observations  have  been  enlisted  to  demonstrate  more  fully, 
if  possible,  the  activity  of  the  different  parts  of  the  epithelial  tubule.  Heiden- 
hain,  by  injections  of  indigo-blue  into  the  blood  stream,  followed  by  rapid 
fixation  of  the  kidney  in  alcohol  at  the  proper  stage  of  elimination,  has 
demonstrated  crystals  of  the  pigment  in  the  renal  epithelial  cells  and  in  the 
lumen  of  the  tubule.  He  concluded  that  these  cells  were  actively  eliminating 
the  pigment  by  a  secretory  process.  This  observation  has  been  questioned. 


DIURETICS  439 

But  Heidenhain's  view  is  strengthened  by  Bowman's  observation  that  in  birds 
crystals  of  uric  acid  are  to  be  seen  in  the  cells  of  the  convoluted  tubules,  and 
in  the  lumen  adjacent. 

Only  traces  of  the  sugars  and  proteins  of  the  blood  are  found  in  normal 
urine,  but  when  either  cane  sugar,  peptone,  or  egg  albumin  is  introduced  into 
the  blood  it  is  rapidly  eliminated  by  the  kidney.  Egg  albumin  is  not  essen- 
tially different  from  the  serum  albumin  of  the  blood,  but  the  serum  albumin 
is  not  excreted.  These  are  both  non-dialyzable  compounds.  Dextrose  and 
urea,  both  readily  dialyzable,  present  the  same  comparison,  i.e.,  urea  is  ex- 
creted, while  dextrose  is  not.  If,  however,  the  percentage  of  this  sugar  is 
high,  0.25  per  cent,  or  more,  it  is  then  eliminated.  The  excretion  of  the 
highly  diffusible  sodium  chloride  bears  a  similar  quantitative  relation  to 
excretion.  If  present  in  the  blood  in  relatively  low  amounts  it  is  not  secreted, 
while  if  the  concentration  is  slightly  greater  it  may  be  quickly  eliminated. 


FIG.  303. — Curve  Taken  by  Renal  Onkometer  Compared  with  that  of  an  Ordinary  Blood- 
pressure  Curve,     a,  Kidney  curve;  6,  blood-pressure  curve.     (Roy.) 

Other  inorganic  salts  present  only  in  traces,  are  meanwhile  rapidly  elimin- 
ated. Even  the  rapid  elimination  of  a  slight  excess  of  water  in  the  blood  can 
scarcely  be  explained  on  purely  physical  grounds.  To  discharge  the  water 
across  the  glomerulus  from  the  blood  to  the  urine  requires  an  expenditure 
of  osmotic  pressure  much  greater  than  that  balanced  by  the  blood  pressure. 
That  is,  the  epithelial  cells  must  do  work,  and  the  energy  is  dependent  on 
metabolism  in  the  cells. 

It  would  seem,  therefore,  that  the  separation  of  urine  in  the  kidney  is  a 
secretory  process  dependent  on  the  protoplasmic  activity  of  the  living  renal 
cells,  that  the  apparent  selective  property  of  the  cells  is  a  manifestation  of 
such  activity,  and  that  even  water  is  actively  secreted. 

Diuretics. — Certain  substances  increase  the  flow  of  urine  and  are 
called  diuretics.  They  act  directly  on  the  renal  epithelium,  for  example 
urea,  or  indirectly  on  the  circulatory  system  to  increase  the  flow  of  blood. 
Digitalis  is  a  well-known  diuretic  which  increases  the  efficiency  of  the  circula- 
tion. It  also  stimulates  the  renal  epithelium  with  the  production  of  a  marked 
increase  in  the  flow  of  urine.  Caffeine  diuresis  can  best  be  explained  on  an 
assumed  stimulating  action  of  the  renal  epithelium.  Urea  introduced  into 


440  EXCRETION 

the  blood  produces  a  copious  secretion  of  urine.  Both  urea  and  the  saline 
diuretics  induce  a  flow  of  urine  out  of  all  proportion  to  the  osmotic  changes 
produced,  and  they  may  be  regarded  as  direct  stimulators  of  the  renal 
epithelium,  a  view  supported  by  their  stimulative  action  on  other  tissues. 

THE  DISCHARGE  OF  THE  URINE. 

As  each  portion  of  urine  is  secreted,  it  propels  that  which  is  already  in  the 
uriniferous  tubes  onward  into  the  pelvis  of  the  kidney.  Thence  it  passes 
through  the  ureter  into  the  bladder,  from  which  at  intervals  it  is  discharged 
to  the  exterior.  The  rate  and  mode  of  entrance  of  urine  into  the  bladder 
has  been  watched  in  cases  of  ectopia  vesicse,  i.e.,  cases  in  which  fissures  in 
the  anterior  or  lower  part  of  the  walls  of  the  abdomen  and  of  the  front  wall 
of  the  bladder  expose  to  view  the  orifices  of  the  ureters.  The  urine  does  not 
enter  the  bladder  at  any  regular  rate,  nor  is  there  a  synchronism  in  its  move- 
ment through  the  two  ureters.  Ordinarily  two  or  three  drops  enter  the 
bladder  every  minute,  each  drop  as  it  enters  first  raising  up  the  little  papilla 
on  which  the  ureter  opens,  and  then  passing  through  the  orifice,  which  at 
once  again  closes  like  a  sphincter.  Its  flow  is  aided  by  the  peristaltic  con- 
tractions of  the  ureters,  and  is  increased  in  deep  inspiration  or  by  straining. 
The  urine  collected  in  the  bladder  is  prevented  from  regurgitation  into  the 
ureters  by  the  mode  in  which  these  pass  through  the  walls  of  the  bladder, 
namely,  by  their  lying  a  half  to  three-quarters  of  an  inch  between  the  muscu- 
lar and  mucous  coats  before  they  turn  rather  abruptly  forward  and  open 
through  the  latter  into  the  interior  of  the  bladder. 

Micturition. — The  contraction  of  the  muscular  walls  of  the  bladder 
may  by  itself  expel  the  urine  with  little  or  no  help  from  other  muscles.  The 
vesicular  pressure  is  increased  in  the  voluntary  act  by  the  contraction  of 
the  abdominal  and  other  expiratory  muscles  which  bear  on  the  abdominal 
viscera,  thus  aiding  in  the  expulsion  of  the  contents  of  the  bladder.  The 
diaphragm  is  at  the  same  time  fixed  in  contraction  and  the  sphincter  of  the 
bladder  relaxes.  The  pressure  within  the  bladder  under  the  combined  con- 
tractions of  these  expulsive  muscles  sometimes  amounts  to  8  to  10  cm.  of 
mercury.  The  act  is  completed  by  the  accelerator  urinae  muscle,  which,  as 
its  name  implies,  quickens  the  stream  and  expels  the  last  drop  of  urine  from 
the  urethra.  The  act  is  under  the  regulative  control  of  a  nervous  center  in 
the  lumbar  spinal  cord,  through  which,  as  in  the  case  of  the  similar  center  for 
defecation,  the  various  muscles  concerned  are  coordinated  in  their  action. 
It  is  well  known  that  the  act  may  be  reflexly  induced,  e.g.,  in  children  who 
suffer  from  intestinal  worms  or  other  such  irritation.  Generally  the  afferent 
impulses  which  set  up  the  reflexes  leading  to  the  desire  to  micturate  are  ex- 
cited by  overdistention  of  the  bladder,  or  sometimes  by  a  few  drops  of  urine 
passing  into  the  urethra.  This  impulse  passes  up  to  the  lumbar  center  or 


STRUCTURE 


441 


centers,  and  reflexly  produces  on  the  one  hand  inhibition  of  the  sphincter 
and  on  the  other  contraction  of  the  muscles  of  the  f  undus  and  other  necessary 
muscles  for  the  expulsion  of  the  contents  of  the  bladder.  In  the  voluntary 
act  these  motor  centers  are  stimulated  to  activity  by  impulses  coming  from 
the  higher  cerebral  centers. 


THE  STRUCTURE  AND  EXCRETORY  FUNCTIONS  OF  THE  SKIN. 

The  skin  serves,  i,  as  an  external  integument  for  the  protection  of  the 
deeper  tissues,  and  2,  as  a  sensitive  organ  in  the  exercise  of  touch,  a  subject 
to  be  considered  in  the  chapter  on  the  Special  Senses.  It  is  also,  3,  an  im- 
portant secretory  and  excretory  organ;  and  4,  an  absorbing  organ.  5,  It 
plays  an  important  part  in  the  regulation  of  the  temperature  of  the  body  by 


-a/ 


FIG.  304. — Vertical  Section  of  the  Epidermis  of  the  Prepuce,  a,  Stratum  corneum, 
of  very  few  layers,  the  stratum  lucidum  and  stratum  granulosum  not  being  distinctly 
represented;  b,  c,  d,  and  e,  the  layers  of  the  stratum  Malpighii,  a  certain  number  of  the  cells 
in  layers,  d;  and  e  showing  signs  of  segmentation;  layer  c  consists  chiefly  of  prickle  or  ridge 
and  furrow  cells;/,  basement  membrane;  g,  cells  in  cuds  vera.  (Cadiat.) 

controlling  the  loss  of  heat,  i.e.,  a  temperature-regulating  function,  discussed 
in  the  chapter  on  Animal  Heat. 

Structure. — The  skin  consists  principally  of  a  vascular  tissue  named 
the  corium,  derma,  or  cutis  vera,  and  of  an  external  covering  of  epithelium 
termed  the  epidermis  or  cuticle.  Within  and  beneath  the  corium  are  em- 
bedded several  organs  with  special  functions,  namely,  sudoriferous  glands, 
cesbaeous  glands,  and  hair  follicles;  and  on  its  surface  are  sensitive  papilla. 


442 


EXCRETION 


The  so-called  appendages  of  the  skin — the  hair  and  nails — are  modifications 
of  the  epidermis. 

The  epidermis  is  composed  of  several  strata  of  cells  of  various  shapes  and 
sizes;  it  closely  resembles  in  its  structure  the  epithelium  of  the  mucous  mem- 
brane that  lines  the  mouth  or  covers  the  cornea.  The  following  four  layers 
may  be  distinguished. 


FIG.  305. — Vertical  Section  of  Skin.  A,  Sebaceous  gland  opening  into  hair  follicle; 
B,  muscular  fibers;  C,  sudoriferous  or  sweat  gland;  D,  subcutaneous  fat;  E,  fundus  of  hair 
follicle,  with  hair-papillae.  (Klein.) 

The  stratum  lucidum,  a  homogeneous  membrane,  consisting  of  squamous 
cells  closely  arranged,  in  some  of  which  a  nucleus  can  be  seen.  Stratum 
granulosum,  consisting  of  one  layer  of  flattened,  fusiform,  distinctly  nucleated 
cells.  Stratum  Malpighii  or  rete  mucosum  consists  of  many  strata  of  cells. 
The  deepest  cells,  placed  immediately  above  the  cutis  vera,  are  columnar 
with  oval  nuclei,  succeeded  by  a  number  of  layers  of  more  or  less  polyhedral 
cells  with  spherical  nuclei;  the  more  superficial  layers  are  considerably 
flattened.  The  deeper  surface  of  the  rete  mucosum  is  accurately  adapted 


GLANDS    OF   THE    SKIN  443 

to  the  papillae  of  the  true  skin,  being,  as  it  were,  molded  on  them.  It 
is  very  constant  in  thickness  in  all  parts  of  the  skin.  The  cells  of  the  middle 
layers  of  the  stratum  Malpighii  are  connected  by  processes,  and  thus  form 
prickle  cells,  figure  28.  The  pigment  of  the  skin,  in  the  deeper  cells  of  the  rete 
mucosum,  causes  the  various  tints  observed  in  different  individuals  and  differ- 
ent races.  The  epidermis  maintains  its  thickness  in  spite  of  the  constant  wear 
and  tear  to  which  it  is  subjected.  The  columnar  cells  of  the  deepest  layer 
of  the  rete  mucosum  elongate,  multiply  by  division,  the  new  cells  produced 
being  pushed  toward  the  free  surface  of  the  skin.  There  is  thus  a  constant 
production  of  fresh  cells  in  the  deeper 
layers,  and  a  constant  throwing  off  of 
old  ones  from  the  free  surface.  When 
these  two  processes  are  accurately 
balanced,  the  epidermis  maintains  its 
thickness.  When  by  intermittent 
pressure  a  more  active  cell  growth  is 
stimulated,  the  production  of  cells  ex- 
ceeds their  waste  and  the  epidermis 
increases  in  thickness,  as  we  see  in 
the  horny  hands  of  the  laborer. 

The  dermis,  or  cutis  vera  or  true  FlG    3o6>_Terminal    Tubules    of 

skin,  is  a  dense  and  tough,  but  yielding     Sudoriferous    Glands,    Cut    in    Various 


and  highly  elastic  structure  support-  "  ** 


ing  the  epidermis.     It  is  composed  of 

areolar  connective  tissue  interwoven  in  all  directions  and  forming  numerous 
spaces  by  its  interlacements.  These  areolae  in  the  deeper  layers  of  the  cutis 
are  usually  filled  with  masses  of  fat  cells,  figure  305.  Unstriped  muscu- 
lar fibers  are  also  abundantly  present,  especially  in  the  skin  of  animals  which 
erect  the  hairs  with  greater  ease  than  is  usually  the  case  with  man. 

There  is  a  rich  network  of  blood  vessels  to  the  dermis.  In  the  dermal 
papillae  and  about  the  sweat  glands  there  are  special  loops  of  capillaries. 
Nerve  fibers  are  also  distributed  to  the  papillae. 

The  special  nerve  terminations  in  the  skin  have  been  described  on  page  75 

Glands  of  the  Skin.  —  The  skin  possesses  glands  of  two  kinds:  Sudor- 
iferous or  sweat  glands,  and  the  sebaceous  or  oil  glands. 

A  sudoriferous  or  sweat  gland  consists  of  a  small  lobular  mass,  formed 
of  a  coil  of  a  simple  tubular  gland,  surrounded  by  blood  vessels,  and  em- 
bedded in  the  subcutaneous  adipose  tissue,  figure  305,  C.  The  duct  as- 
cends from  this  coiled  mass  for  a  short  distance  in  a  spiral  manner  through 
the  cutis  and  the  epidermis,  and  then  opens  on  the  surface  of  the  skin.  In 
the  parts  where  the  epidermis  is  thin,  the  ducts  themselves  are  thinner 
and  more  nearly  straight  in  their  course. 

The  duct  is  lined  with  a  layer  of  columnar  epithelium  continuous  with 


444 


EXCRETION 


the  epidermis.  The  coiled  or  secreting  portion  of  the  gland  is  lined  with 
at  least  two  layers  of  short  columnar  cells  with  very  distinct  nuclei,  figure  306. 
The  lumen  is  distinctly  bounded  by  a  special  lining  of  cuticle. 

The  sudoriferous  glands  are  abundantly  distributed  over  the  whole  sur- 
face of  the  body;  but  are  especially  numerous,  as  well  as  very  large,  in  the 
skin  of  the  palm  of  the  hand  and  of  the  sole  of  the  foot.  The  glands  by 
which  the  peculiarly  odorous  matter  of  the  axillae  and  groin  is  secreted  form 

a  nearly  complete  layer  under  the  cutis, 
and  are  like  the  ordinary  sudoriferous 
glands,  except  in  being  larger  and  having 
very  short  ducts. 

The  peculiar  bitter  yellow  substance 
secreted  by  the  skin  of  the  external 
auditory  passage  is  named  cerumen,  and 
the  glands  themselves  ceruminous  glands; 
but  they  do  not  much  differ  in  structure 
from  the  ordinary  sudoriferous  glands. 

The  sebaceous  glands,  figures  305 
and  306,  like  sudoriferous  glands,  are 
abundant  in  most  parts  of  the  surface  of 
the  body,  particularly  in  parts  largely 
supplied  with  hair,  as  the  scalp  and  face. 
They  are  thickly  distributed  about  the 
entrance  of  the  various  passages  into  the 
body,  as  the  anus,  nose,  lips,  and  ex- 
ternal ear.  They  are  entirely  absent 
from  the  palmar  surface  of  the  hand  and 
the  plantar  of  the  foot.  They  are 
racemose  glands  composed  of  an  aggre- 
gate of  small  tubes  or  sacculi  lined  with  columnar  epithelium  and  filled 
with  an  opaque  white  substance,  like  soft  ointment,  which  consists  of 
broken-up  epithelial  cells  which  have  undergone  fatty  degeneration.  Mi- 
nute capillary  vessels  overspread  them;  and  their  ducts  open  on  either  the 
surface  of  the  skin,  close  to  the  hair,  or,  which  is  more  usual,  directly  into 
the  follicle  of  the  hair.  In  the  latter  case,  there  are  generally  two  or  more 
glands  to  each  hair,  figure  306. 

The  story  of  the  structure  and  development  of  such  epithelial  structures 
as  the  hair  and  nails  is  best  left  to  the  histologist,  to  whom  the  student  is 
referred. 

The  Excretory  Function  of  the  Skin. — The  function  of  the  skin  which 
is  of  special  interest  to  this  chapter  is  that  of  the  secretion  of  the  sweat.  The 
fluid  secreted  by  the  sweat  glands  is  usually  formed  so  gradually  that  the 
watery  portion  of  it  escapes  by  evaporation  as  fast  as  it  reaches  the  surface. 


FIG.  307. — Sebaceous  Gland  from 
Human  Skin.  (Klein  and  Noble 
Smith.) 


THE   EXCRETORY   FUNCTION    OF   THE    SKIN  445 

But  during  strong  exercise,  exposure  to  great  external  warmth,  in  some 
diseases,  and  when  evaporation  is  prevented,  the  secretion  becomes  more 
sensible  and  collects  on  the  skin  in  the  form  of  drops  of  fluid. 

The  -perspiration,  as  the  term  is  sometimes  employed  in  physiology,  in- 
cludes all  that  portion  of  the  secretions  and  exudations  from  the  skin  which 
are  thrown  on  the  surface  by  the  sweat  glands.  As  a  matter  of  fact,  this  is 
mingled  with  various  substances  lying  on  the  surface  of  the  skin.  The  con- 
tents of  the  sweat  are,  in  part,  matters  capable  of  assuming  the  form  of  vapor, 
such  as  carbonic  acid  and  water,  and  in  part  other  matters  which  are 
deposited  on  the  skin,  and  mixed  with  the  sebaceous  secretions. 

The  secretion  of  the  sebaceous  glands  and  hair  follicles  consists  of  cast- 
off  epithelium  cells,  with  nuclei  and  granules,  together  with  an  oily  material 
and  extractive  matter.  In  certain  parts,  also,  it  is  mixed  with  a  peculiar 
odorous  principle,  which  contains  caproic,  butyric,  and  other  fatty  acids. 
It  is  similar  in  composition  to  the  unctuous  coating,  or  vernix  caseosa,  which 
is  formed  on  the  body  of  the  fetus  while  in  the  uterus,  and  which  contains 
ordinary  fat  and  cholesterol  esters  with  fatty  acids.  This  sebaceous  secre- 
tion serves  the  purpose  of  keeping  the  skin  moist  and  supple,  and,  by  its  oily 
nature,  of  both  hindering  the  evaporation  from  the  surface  and  guarding 
the  skin  from  the  effects  of  the  long-continued  action  of  moisture.  But 
while  it  thus  serves  local  purposes,  its  removal  from  the  body  entitles  it  to  be 
listed  among  the  excretions  of  the  skin. 

CHEMICAL  COMPOSITION  OF  SWEAT. 

Water 995 

Solids:   5 

Organic  acids   (formic,   acetic,  butyric,   pro- 

pionic,  caproic,  caprylic) 0.9 

Salts,  chiefly  sodium  chloride 1.8 

Neutral  fats  and  cholesterol 0.7 

Extractives  (including  urea),  with  epithelium   i  .6 


The  sweat  is  a  colorless,  slightly  turbid  fluid,  alkaline,  neutral  or  acid  in 
reaction,  of  a  saltish  taste,  and  peculiar  characteristic  odor. 

Of  the  several  substances  it  contains,  however,  only  the  carbon  dioxide 
and  water  need  particular  consideration. 

The  quantity  of  water  vapor  excreted  from  the  skin  is,  on  an  average, 
between  750  and  1,000  c.c.  daily.  This  subject  has  been  very  carefully 
investigated  by  Lavoisier  and  Sequin.  The  latter  chemist  enclosed  his 
body  in  an  air-tight  bag  provided  with  a  mouthpiece.  The  bag  was  closed 
by  a  strong  band  above,  and  the  mouthpiece  adjusted  and  gummed  to  the 
skin  around  the  mouth.  He  was  weighed,  then  remained  quiet  for  several 


446  EXCRETION 

hours,  after  which  time  he  was  again  weighed.  The  difference  in  the  two 
weights  indicated  the  amount  of  loss  by  pulmonary  exhalation.  Having 
taken  off  the  air-tight  dress,  he  was  immediately  weighed  again,  and  a  fourth 
time  after  a  certain  interval.  The  difference  between  the  two  weights  last 
ascertained  gave  the  amount  of  the  cutaneous  and  pulmonary  exhalation 
together;  by  subtracting  from  this  the  loss  by  pulmonary  exhalation  alone, 
while  he  was  in  the  air-tight  dress,  he  ascertained  the  amount  of  cutaneous 
transpiration.  The  average  loss  by  cutaneous  and  pulmonary  exhalation  in 
a  minute  during  a  state  of  rest  was  eighteen  grains — the  minimum  eleven 
grains,  the  maximum  thirty-two  grains.  Of  the  eighteen  grains,  eleven 
passed  off  by  the  skin  and  seven  by  the  lungs. 

The  quantity  of  watery  vapor  lost  by  transpiration  is,  of  course,  influenced 
by  all  external  circumstances  which  affect  the  exhalation  from  evaporating 
surfaces,  such  as  the  temperature,  the  hygrometric  state,  and  the  stillness 
of  the  atmosphere.  But,  of  the  variations  to  which  it  is  subject  under  the 
influence  of  these  conditions,  no  calculation  has  been  exactly  made. 

The  quantity  of  carbon  dioxide  exhaled  by  the  skin  on  an  average  is 
said  to  be  about  one  two-hundredth  of  that  eliminated  by  the  pulmonary 
respiration. 

The  cutaneous  exhalation  is  most  abundant  in  the  lower  classes  of  ani- 
mals, more  particularly  the  naked  amphibia,  as  frogs  and  toads,  whose  skins 
are  thin  and  moist,  and  readily  permit  an  interchange  of  gases  between  the 
circulating  blood  and  the  surrounding  atmosphere.  Bischoff  found  that, 
after  the  lungs  of  frogs  had  been  tied  and  cut  out,  from  3  to  4  c.c.  of  carbon 
dioxide  gas  was  exhaled  by  the  skin  in  eight  hours.  And  this  quantity 
is  very  large,  when  it  is  remembered  that  a  full-sized  frog  will  generate  only 
about  10  c.c.  of  carbon  dioxide  by  his  lungs  and  skin  together  in  six  hours. 

The  importance  of  the  respiratory  function  of  the  skin,  which  was  once 
thought  to  be  proved  by  the  speedy  death  of  animals  whose  skins,  after  re- 
moval of  the  hair,  were  covered  with  an  impermeable  varnish,  has  been  shown 
by  further  observations  to  have  no  foundation  in  fact.  The  immediate  cause 
of  death  in  such  cases  is  the  interference  with  temperature  regulation. 

Influence  of  the  Nervous  System  on  Sweat  Secretion. — The  secre- 
tion of  sweat  is  closely  connected  with  the  quantity  of  blood  flowing  through 
the  cutaneous  vessels.  The  quantity  of  sweat  increases  with  vaso-dilatation 
and  diminishes  with  vaso-constriction.  The  sweat  glands  are  also  under  the 
control  of  efferent  impulses  passing  to  them  from  the  special  sweat  centers 
in  the  brain  and  spinal  cord  through  special  sweat  nerves.  Thus,  if  the 
sciatic  nerve  be  divided  in  a  cat  and  the  peripheral  end  be  stimulated,  beads  of 
sweat  are  seen  to  appear  upon  the  pad  of  the  corresponding  foot.  The  sweat 
appears  even  though  at  the  same  time  the  blood  vessels  are  constricted,  or 
the  blood  flow  entirely  stopped  by  compression  of  the  aorta,  whereas  if  atropin 
is  injected  previously  to  the  stimulation,  no  sweat  appears,  although  dila- 


THE    PILO-MOTOR   NERVES    TO    THE    SKIN  447 

tation  of  the  vessels  may  be  present.  Secretion  of  sweat,  too,  may  be  brought 
about  reflexly. 

The  circulation  of  venous  blood  in  the  spinal  bulb  causes  the  sweating  of 
phthisis  and  of  dyspnea  generally,  by  stimulating  the  sweat  center.  If  the 
cat  whose  sciatic  nerve  is  divided  be  rendered  dyspneic,  abundant  sweat 
occurs  upon  the  foot  of  the  uninjured,  and  none  on  the  injured,  side.  The 
effect  of  heat  in  producing  sweating  may  be  both  local  and  general,  and,  again, 
the  various  drugs  which  produce  an  increased  secretion  of  sweat  do  not  all 
act  in  the  same  way;  thus,  there  is  reason  for  thinking  that  pilocarpine  acts 
upon  the  local  apparatus,  that  strychnine  and  picrotoxin  act  upon  the  sweat 
centers,  and  that  nicotine  acts  both  upon  the  central  and  upon  the  local 
apparatus. 

The  special  sweat  nerves  belong  to  the  thoracid  autonomies  or  sym- 
pathetic nerves,  and  issue  from  the  spinal  cord  with  this  outflow,  page 
636.  The  fibers  join  the  chain  ganglia  by  way  of  the  white  rami  and 
are  thence  distributed  to  all  parts  of  the  body.  In  the  case  of  the  hind 
limb  of  the  cat  the  sweat  nerves  arise  from  the  cord  by  the  last  two  or 
three  dorsal  and  first  two  to  four  lumbar  nerves,  pass  to  the  sympathetic 
chain,  and  from  thence  to  the  sciatic  nerve,  following  the  general  course  of 
the  autonomic  nerves  for  this  region.  In  the  case  of  the  fore  limb,  the 
nerves  leave  the  cord  by  the  first  to  the  sixth  dorsal  roots  pass  into  the 
thoracic  sympathetic  and  then  join  the  brachial  plexus,  reaching  the  arm 
through  the  median  and  ulnar  nerves. 

The  Pilo-motor  Nerves  to  the  Skin. — Small  groups  of  smooth  mus- 
cle fibers  are  found  generally  distributed  in  the  skin.  They  are  attached 
to  the  hair  follicles  and  by  their  contractions  produce  erection  of  the  hairs 
or  in  man  the  roughness  called  goose-flesh.  These  muscles  are  innervated 
by  nerve  fibers  that  belong  to  the  thoracic  autonomies.  The  origin  and 
path  of  distribution  corresponds  closely  with  that  of  the  sweat  nerves. 

It  will  be  as  well  to  restate  here  the  other  functions  which  the  skin  sub- 
serves. In  addition  to  its  excretory  office,  we  have  seen  that  it  acts  as  a 
channel  for  absorption.  It  is  also  concerned  with  the  special  senses,  that  of 
touch  and  temperature,  to  the  consideration  of  which  as  well  as  to  its  function 
of  regulating  the  temperature  of  the  body  we  shall  presently  return.  By  its 
general  impermeability  it  prevents  the  loss  of  moisture  of  the  body  by  direct 
evaporation  from  the  tissues.  It  should  be  recollected,  however,  that  apart 
from  these  special  functions,  by  means  of  its  toughness,  flexibility,  and  elastic- 
ity, the  skin  is  eminently  qualified  to  serve  as  the  general  integument  of  the 
body,  for  defending  the  internal  parts  from  external  violence,  while  readily 
yielding  and  adapting  itself  to  their  various  movements  and  changes  of 
position. 


448  EXCRETION 

LABORATORY  EXPERIMENTS  IN  EXCRETION. 
PHYSIOLOGICAL  REACTIONS. 

i.  The  Relation  of  Blood  Flow  through  the  Kidney  to  the  Secre- 
tion of  Urine. — Properly  to  check  this  experiment  one  should  make  three 
determinations:  i,  the  general  blood  pressure;  2,  the  volume  of  the  kid- 
ney; 3,  the  amount  of  urine  secreted.  Anesthetize  a  dog  and  arrange  the 
apparatus  for  taking  the  blood  pressure  as  directed  in  experiment  19.  Pre- 
pare a  renal  onkometer,  see  figures  301  and  302,  and  an  onkograph  for  re- 
cording the  variations  in  the  volume  of  the  kidney.  The  renal  onkometer 
consists  of  a  double  metal  box  to  fit  the  form  of  a  kidney.  The  inner  halves 
of  this  box  should  be  covered  so  loosely  with  very  thin  sheet  rubber  that  the 
rubber  can  be  fitted  into  the  bottom  of  the  cup  without  undue  tension.  The 
rubber  must  be  sealed  to  the  outer  edges  of  this  inner  cup  with  rubber  cement 
and  allowed  to  dry.  When  it  is  completely  dried  the  inner  cup  should  be  ad- 
justed to  the  outer,  and  the  spaces  enclosed  by  the  rubber  sheet  filled  with 
water.  Or  the  onkometer  may  be  closed  with  parchment  and  filled  with  oil 
as  described  in  experiment  23  on  the  Circulation.  The  half  of  the  onkom- 
eter that  comes  against  the  wall  of  the  body  cavity  of  the  animal  should  be 
completely  closed  with  a  stopper  before  the  instrument  is  adjusted  to  the 
kidney.  Now  adjust  the  onkometer  to  the  kidney,  taking  care  to  place 
the  renal  arteries,  veins,  and  ureter  in  the  tube  in  such  a  way  as  not  to  com- 
press them.  Fill  the  outer  cup  with  water  and  connect  this  cavity  by  a 
two-way  cannula  with  the  recording  onkograph.  In  practice  it  is  more 
satisfactory  if  one  introduces  between  the  onkometer  and  onkograph  an  over- 
flow bottle  or  bulb,  adjusted  to  maintain  the  constant  pressure  on  the  kidney. 
This  direction  varies  from  the  usual  one  in  that  rubber  sheeting  instead  of 
parchment  is  used  to  cover  the  inner  cup  of  the  onkometer,  a  method  that 
permits  the  use  of  water  instead  of  oil.  Recently  Jackson  has  introduced 
an  air  onkometer  that  is  simpler  to  prepare  and  adjust.  The  seal  is  se- 
cured by  wrapping  it  in  folds  of  omentum  after  inserting  the  kidney.  An 
air  recorder  must  of  course  be  used. 

Isolate  and  insert  a  small  cannula  into  the  ureter.  This  cannula  should 
be  clamped  in  a  stand  at  a  level  as  little  above  that  of  the  kidney  as  possible. 
The  urine  secreted  may  be  collected  in  a  loc.c.  graduated  cylinder  and 
measured  at  intervals  of  5  or  10  minutes.  Or,  if  the  outflow  is  scanty,  it  may 
be  allowed  to  drop  on  a  tambour  recording  apparatus,  the  rate  of  dropping 
being  indicative  of  the  rapidity  of  secretion. 

Determine  the  normal  rate  of  secretion  of  a  dog  under  constant  anesthesia. 
The  anesthesia  should  be  medium  to  light,  but  should  be  kept  very  uni- 
form so  as  to  maintain  a  strong  blood  pressure.  Note  the  effect  on  secre- 
tion and  the  corresponding  effect  on  blood  pressure  and  the  kidney  volume 
produced  by  vagus  inhibitions.  Section  the  vagus  nerves  and  produce  in- 


REACTION  449 

hibJtion  by  stimulating  the  peripheral  end  of  the  vagus.  In  this  instance 
there  are  no  reflexes  to  complicate  the  experiment,  so  that  the  fall  in  blood 
pressure  is  a  direct  cardiac  effect.  Stimulate  the  central  end  of  the  vagus 
which  produces  a  fall  of  blood  pressure  through  the  vaso-motor  system. 
There  should  be  a  normal  period  of  at  least  ten  minutes  following  each  ex- 
periment to  allow  the  secretion  of  the  kidney  to  return  to  the  normal. 

Expose  the  splanchnic  nerves  at  the  point  where  they  pass  beneath  the 
diaphragm  into  the  abdominal  cavity.  Adjust  a  pair  of  shielded  electrodes, 
close  the  cavity,  and,  when  the  animal  has  returned  to  the  normal  uniform 
rate  of  secretion  and  of  blood  pressure,  stimulate  the  splanchnic  nerves. 
The  splanchnics  contain  vaso-constrictor  nerves  for  the  kidney.  The  on- 
kometer  experiment  should,  therefore,  demonstrate  a  sharp  decrease  in  the 
volume  of  the  organ,  while  the  blood  pressure  is  only  slighty  changed. 
Inject  10  cubic  centimeters  of  5  per  cent,  potassium  nitrate  or  other  inor- 
ganic diuretics  intravenously.  These  tests  should  be  repeated  with  long 
intervals  for  readjustment. 

In  this  connection  demonstrate  the  influence  of  deep  chloroform  anesthesia 
on  urinary  secretion.  The  chloroform  should  be  pushed  to  the  danger  limit 
and  maintained  there  for  a  couple  of  minutes  or  more.  Compare  the  rapidity 
of  the  recovery  of  blood  pressure  with  the  recovery  of  the  rate  of  secretion. 

2.  Secretory  Nerves  for  the  Sweat  Glands. — Langley  has  mapped 
out  the  paths  of  the  secretory  nerves  for  the  sweat  glands.     He  has  shown 
that  in  the  cat  these  fibers  are  distributed  to  the  hind  limb  through  the  sciatic. 
Anesthetize  a  half-grown  cat,  isolate  the  sciatic  nerve,  cut  it  and  stimulate  the 
peripheral  end  with  a  medium  to  strong  induction  current.     After  a  few 
moments  beads  of  perspiration  will  appear  on  the  pads  of  the  foot,  which 
should  therefore  be  carefully  examined  before  the  experiment. 

URINE  ANALYSIS. 

3.  Daily  Quantity. — Determine  the  total  quantity,  for  24  hours,  of 
urine  secreted  through  a  period  of  3  or  4  days,  beginning  and  ending  the 
period  at  a  definite  hour  in  the  day,  preferably  on  rising  in  the  morning. 
The  daily  secretion  varies  through  wide  extremes,  depending  upon  the  quan- 
tity of  liquid  taken  in  the  food,  the  daily  exercise,  the  temperature,  etc.,  etc. 
In  the  analysis  of  urine  it  is  always  better  to  take  a  mixed  24-hour  sample. 

4.  Specific  Gravity. — Determine  the  specific  gravity  of  24-hour  urine. 
This  is  done  by  the  instrument  known  as  the  urinometer  which  carries 
a  graduated  scale  at  the  neck.     Care  should  be  taken  to  float  the  urinom- 
eter so  that  it  does  not  come  in  contact  with  the  measuring  cylinder.     The 
scale  should  be  read  at  the  bottom  of  the  meniscus. 

5.  Reaction. — Determine  the  reaction  of  perfectly  fresh  urine,  using 
litmus  paper.     The  normal  urine  is  slightly  acid  under  ordinary  conditions, 
due  to  the  presence  of  acid  phosphates  or  perhaps  in  some  cases  to  traces  of 
free  organic  acid. 


450 


EXCRETION 


After  standing  some  time  the  reaction  is  usually  alkaline,  owing  to  fer- 
mentation processes.  The  reaction  may  vary  also  according  to  the  food, 
vegetable  foods  tending  to  produce  alkaline  urine,  while  with  animal  foods 
the  reaction  is  acid. 

6.  The  Total   Quantity   of   Solids. — Determine   the  solids   of  urine 
by  evaporating  25  cc.  of  a  mixed  sample  of  urine  to  dryness  in  a  weighed 
platinum  or  porcelain  dish  over  a  water  bath.     The  residue  should  be  dried 
to  constant  weight  in  a  drying  oven  at  105°  C. 

A  useful  rule  for  approximately  estimating  the  total  solids  in  any  given 
specimen  of  healthy  urine  is  to  multiply  the  last  two  figures  representing  the 
specific  gravity  by  2 . 33.  Thus,  in  urine  of  specific  gravity  1025,  2.33X25  = 
58.25  grains  of  solids  are  contained  in  1,000  grains  of  the  urine.  Or  the 
total  solids  are  5.825  per  cent.  In  using  this  method  it  must  be  remem- 
bered that  the  limits  of  error  are  much  wider  in  diseased  than  in  healthy 
urine. 

The  solids  of  urine  consist  of  inorganic  salts  of  sodium,  potassium,  and 
calcium,  and  of  a  long  list  of  organic  compounds,  chiefly  nitrogenous. 

7.  Chlorides. — Large  quantities  of  sodium  chloride  are  always  present 
in  the  normal  urine.     Add  ammonia  to  25  or  50  cc.  of  albumin-free  urine 

and  heat  to  precipitate  earthy  phosphates,  filter.  To  a 
sample  of  the  filtrate  add  an  excess  of  strong  nitric  acid 
and  a  few  drops  of  i  per  cent,  silver  nitrate.  A  white 
flocculent  precipitate  of  silver  chloride  comes  down.  This 
precipitate  is  soluble  in  an  excess  of  ammonia.  Repre- 
cipitate  by  adding  nitric  acid  again.  The  test  may  be 
made  without  removing  the  phosphates,  though  in  this 
case,  upon  adding  ammonia,  the  disappearance  of  the 
silver  precipitate  is  complicated  by  the  appearance  of 
insoluble  phosphates. 

The  chlorides  may  be  estimated  quantitatively  by 
Volhard's  method,  or  some  one  of  its  modifications,  which 
depends  upon  the  determination  of  the  amount  of  chloride 
precipitated  by  the  silver.  The  student  is  referred  to  chem- 
ical text-books  for  this  and  other  quantitative  methods. 

8.  Sulphates. — Sulphates  exist  in  the  urine  both  in 
inorganic  and  organic  compounds,  chiefly  the  former. 
Add  a  few  drops  of  hydrochloric  acid  to  a  sample  of 
urine  in  a  test-tube,  then  a  solution  of  barium  chloride,  the 
insoluble  barium  sulphate  settles  out.  If  the  test  is  made  on  the  normal 
urine  without  the  addition  of  the  acid,  the  inorganic  sulphate  will  be  pre- 
cipitated, while  the  ethereal  or  compound  sulphate  will  remain  in  solution 
and  can  be  filtered  off.  This  filtrate,  when  boiled  with  strong  hydrochloric 
acid  to  10  per  cent,  over  a  water  bath  for  a  short  time,  will  have  the  sul- 


FIG.  308.— The 
Urinometer. 


URIC   ACID 


451 


phates  split  off  from  the  organic  radicle  and  may  be  precipitated  by  the 
addition  of  barium  chloride  in  hot  solution. 

9.  Phosphates. — The  phosphates  of  urine  consist  of  the  earthy  and 
alkaline  salts,  the  latter  predominating.  Take  a  50  cc.  sample  of  urine, 
add  strong  ammonia,  and  heat.  The  phosphates  of  calcium  and  magnesium 
separate  out,  as  they  are  insoluble  in  alkaline  solution.  Filter. 

To  the  nitrate  add  a  solution  of  magnesium  sulphate.  This  precipitates 
the  sodium  and  potassium  phosphates  as  a  triple  phosphate  of  magnesium 
which  is  insoluble.  Test  for  phosphates  in  general  are: 

Add  nitric  acid  to  a  sample  of  urine,  warm  gently,  then  add  a  few  drops 
of  10  per  cent,  ammonium  molybdate;  a  yellow  precipitate  of  ammonium 
phospho-molybdate  is  formed.  Or,  add  acetic  acid,  then  a  few  drops  of 
uranium  acetate;  a  bright  yellow  precipitate  of  uranium  ammonium  phos- 
phate is  formed.  These  two  reactions  are  used  as  the  basis 
for  a  quantitative  determination  of  phosphorus. 

10.  The  Preparation  of  Urea. — Take  100  cc.  of  normal 
urine,  evaporate  to  one-half  its  quantity,  and  precipitate  the 
phosphates  and  sulphates  by  adding  a  mixed  solution  of  barium 
hydrate  and  nitrate.  Filter,  evaporate  the  nitrate  to  dryness, 

take  up  in  warm    95   per 
cent,  alcohol,  and  refilter. 
Crystals  of  urea    separate 
out    when    the     alcohol     is 
evaporated  off. 

Evaporate  a  large  sample,  200 
cc.,  of  urine  to  a  syrupy  mass,  add  nitric 
acid.     Crystals   of  urea  nitrate  are  formed. 
Wash  the  crystals  in  dilute  nitric  acid,  then  dis- 
solve in  water.    The  urea  is  set  free  by  adding  bar- 
ium carbonate  until  the  carbon  dioxide  ceases  to 
come  off.     Filter,  evaporate  over  a  water  bath  to  dry- 
ness,  and  dissolve  the  urea  in  95  per  cent,  alcohol ;  de- 
cant, and  recrystallize  by  evaporating  off  the  alcohol. 

11.  Urea  Determination  by  Doremus'  Ureometer.— Fill  the  ureom- 
eter  with   hypobromite  of  sodium  solution.     Take  a  sample  of  urine  in 
the  pipet  which   accompanies  the  instrument,   drawing  it  exactly  to  the 
mark.     Insert  the  pipet  past  the  bend  of  the  ureometer  and  slowly  and 
carefully  empty  the  urine  so  as  not  to  lose  any  of  the  liberated  nitrogen. 
The  instrument  is  graduated  to  read  off  the  percentage  of  urea  directly. 

12.  Uric  Acid.— Concentrate   over   a  water  bath  500  cc.  of  urine  to 
100  cc.  and  boil  with   10  cc.  or  more  of  strong  hydrochloric  acid.     Upon 
cooling,  crystals  of  uric  acid  are  formed.     Decant  the  supernatant  liquid  and 
wash  the  crystals  with  a  few  cubic  centimeters  of  10  per  cent,  hydrochloric 
acid.     Dissolve  the  crystals  and  test. 


FIG.  309. — Doremus' 
Ureometer. 


452  EXCRETION 

The  Mwexide  Test. — Add  to  2  cc.  of  uric  acid  solution  in  a  test-tube  an 
equal  quantity  of  nitric  acid.  Heat  gently,  a  reddish  ring  forms  at  the  point 
of  contact  between  the  nitric  acid  and  uric  acid  solution.  Cool  and  add 
ammonia  carefully.  The  color  ring  deepens  to  a  purple  color.  This  test 
succeeds  well  by  evaporating  a  few  drops  of  uric  acid  on  a  porcelain  plate. 
Add  to  the  stain  a  drop  of  concentrated  nitric  acid  and  evaporate.  Concen- 
tric rings  of  reddish  color  will  be  formed.  This  color  deepens  to  reddish- 
purple  when  a  drop  of  ammonia  is  added. 

13.  Creatinin. — Test    20    cc.   of  urine  in  a  beaker  for  creatinin  by 
adding  a  cubic  centimeter  of  dilute  solution  of  sodium  nitroprusside  and  then 
weak  sodium  hydrate.     A  ruby-red  color,  which  quickly  turns  yellow,  indi- 
cates the  presence  of  creatinin  (Weyl's  reaction).     If  the  yellow  solution 
has  an  excess  of  acetic  acid  added  and  is  then  boiled,  it  turns  first  green  and 
later  blue,  forming  ultimately  a  precipitate  of  Prussian  blue.     Urine  mixed 
with  picric  acid  gives  a  red  coloration  when  made  alkaline  with  caustic 
alkali  solution. 

14.  Total  Nitrogen  in  Urine. — Determine  the  total  nitrogen  in  a  sample 
of  urine  by  the  Kjeldahl  method.     This  method  depends  upon  the  con- 
version of  all  the  nitrogen  to  ammonia,  the  distillation  of  this  ammonia 
into  a  known  quantity  of  sulphuric  acid,  and  the  final  titration  of  the  excess 
of  sulphuric  acid  when  the  distillation  is  complete.     The  computation  is 
made  on  the  basis  that  i  cc.  of  a  normal  sulphuric  acid  is  equivalent  to 
i    cc.    normal   sodium   hydrate,  and  that  in  turn  to  i  cc.  of  ammonium 
hydrate.     The  ammonia  neutralizes  a  portion  of  the  sulphuric  acid  in  the 
distillation.     One  cc.  of  normal  ammonium  hydrate  contains  0.014  gram 
nitrogen,  from  which  the  total  nitrogen  in  the  sample  used  can  be  readily 
computed. 

15.  Pigments  of  Urine. — The  normal  color  of  the  urine  is  due  to  the 
presence  of  a  pigment,  urobilin.     Prepare  urobilin  by  adding  lead  acetate 
to  a  200  cc.  sample  of  urine.     A  precipitate  forms  which  carries  down  the 
coloring  mater.     Filter.     Add  acid  alcohol  to  the  precipitate  to  extract  the 
coloring  mater,  refilter,  which  gives  a  deep  yellow  solution.     Shake  up  with 
a  few  cubic  centimeters  of  chloroform  which  dissolves  the  pigment.     Draw 
off  the  chloroform  solution  and  allow  to  evaporate.     The  residue  is  a  brown- 
ish mass  of  urobilin. 

1 6.  Test  for  Indican,  Obermayer's. — Take  in  a  test  tube  i  cc.  Ober- 
mayer's  reagent  (.2  to  .4  per  cent,  ferric  chloride  in  concentrated  hydro- 
chloric acid)  and  an  equal  part  urine  to  be  tested,  then  add  2  cc.  of  chloro- 
form. Thoroughly  shake.  A  blue  color  will  develop  if  indican  be  present. 

ABNORMAL  CONSTITUENTS  OF  URINE. 

Many  abnormal  constituents  may  appear  in  the  urine  under  pathological 
conditions,  only  two  of  which  will  be  mentioned  here. 
28 


QUANTITATIVE    DETERMINATION    OF    SUGAR   IN    THE    URINE    453 

17.  Albumin  in  the  Urine. — The  detection  of  the  presence  of  albumin, 
albuminuria,  is  of  considerable  clinical  importance.     The  following  are  the 
standard  tests  which  present  no  special  difficulty  except  when  traces  only 
are  present. 

Heat  Coagulation. — Take  a  half  test-tube  of  urine,  boil,  and  add  a  drop 
of  dilute  acetic  acid.  A  white  coagulum  indicates  the  presence  of  albumin. 
A  faint  cloudy  appearance  indicates  traces. 

Nitric  Acid  Test. — To  5  cc.  of  strong  nitric  acid  in  a  conical  test-tube 
add  10  or  15  c.c.  of  urine,  pouring  it  gently  down  the  inclined  side  of  the  glass. 
Allow  the  glass  to  stand  for  a  few  minutes,  when  a  white  coagulum  appears 
just  above  the  line  of  contact  of  the  acid  with  the  urine.  This  test,  known 
as  Heller's  test,  will  usually  indicate  the  presence  of  traces  of  albumin. 

Picric  Acid  Test. — Add  picric  acid  to  a  sample  of  urine.  A  whitish  pre- 
cipitate of  albumin  will  appear  at  the  line  of  contact,  as  in  the  preceding  test. 

Citric  acid  two  parts  and  picric  acid  one  part,  when  boiled  with  urine 
will  coagulate  minute  traces  of  the  protein. 

1 8.  Detection  of  Sugar  in  the  Urine. — F 'ehling's  Test. — The  pres- 
ence of  sugar  in  the  urine  can  usually  be  detected  by  Trommer's  test, 
which  depends  upon  the  reduction  of  copper  sulphate  in  the  presence  of 
strong  alkali.     Boil  fresh  Fehling's  solution  and  add  to  it  a  few  drops  of 
urine.     When  sugar  is  present  a  reddish-yellow  precipitate  of  copper  oxide 
comes  out.     The  test  should  be  set  away  for  a  few  minutes  when,  if  only 
traces  of  the  reduction  are  present,  a  reddish-brown  stain  will  appear  on 
the  bottom  of  the  test-tube.     Uric  acid,  if  present  in  excess,  may  produce 
a  slight  precipitation  of  the  copper. 

Fermentation  Test. — If  sugars  are  present  in  the  urine,  they  can  be  de- 
tected by  adding  yeast  to  a  fermentation  tube  filled  with  urine,  the  liberation 
of  carbon  dioxide  indicating  the  presence  of  sugar.  Cane  sugar  does  not 
support  the  growth  of  yeast,  so  it  forms  an  exception  by  this  test. 

Phenyl-Hydrazin  Test. — Phenyl-hydrazin  forms  crystals  of  phenyl- 
glucosazone.  To  10  cc.  of  urine  in  a  small  beaker  add  o.i  of  a  gram  of 
phenyl-hydrazin  hydrochloride  and  a  double  quantity  of  sodium  acetate. 
Heat  in  the  water  bath  for  20  minutes.  Upon  cooling  a  deposit  of  yellow 
crystals  of  phenyl-glucosazone  takes  place  if  glucose  is  present. 

19.  Quantitative  Determination  of  Sugar  in  the  Urine. — Fill  a  10 
cc.  graduated  pipet  with  freshly  prepared  Fehling's  solution.     Take  10  cc. 
of  urine,  measured  with  a  dropping-pipet  into  a  small  beaker,  and  boil. 
While  continuing  to  boil,  add  Fehling's  solution  slowly  and  cautiously  so 
long  as  the  color  is  discharged.     The  amount  of  Fehling's  required  to  reduce 
the  sugar  is  a  measure  of  the  quantity  of  reducing  sugar  present — i  cc.  of 
Fehling's  being  equivalent  to  5  milligrams  of  dextrose. 

For  the  presence  of  blood  pigments  and  other  abnormal  constituents  of 
the  urine,  the  student  is  referred  to  special  handbooks  on  the  subject. 


CHAPTER  XI 
METABOLISM,  NUTRITION,  AND  DIET. 

THE  term  metabolism  means,  literally,  an  exchange  of  material.  In  its 
broadest  physiological  sense  it  includes  the  study  of  the  exchange  of  material 
between  the  living  tissues  of  the  body  and  their  surrounding  media.  This 
includes  the  study  of  the  income  and  outgo  of  material;  the  storing  of  energy- 
yielding  materials  in  the  body;  the  transfer  of  this  potential  energy  into  kinetic 
energy;  and  the  nutritional  processes  within  the  various  tissues.  The  build- 
ing up  of  absorbed  food  material  into  the  protoplasm  of  the  cell  or  of  simpler 
compounds  into  more  complex  ones,  which  may  be  stored  in  the  cell,  is  known 
as  anabolism,  and  the  compounds  themselves  as  anabolites.  The  breaking 
down  of  these  substances  into  simpler  forms,  whereby  the  potential  energy 
of  the  anabolites  is  transformed  into  kinetic  energy,  is  known  as  katabolism, 
and  its  products  as  katabolites. 

In  order  to  form  an  estimate  of  these  processes  going  on  in  the  body,  the 
amount  and  nature  of  the  ingested  material  must  be  known,  as  well  as  the 
amount  of  refuse  or  unused  material  that  passes  out  of  the  alimentary  canal 
as  feces,  and  the  amount  of  excreted  material  from  the  various  excretory 
organs.  It  is  also  necessary  to  know  the  potential  energy  of  the  ingested 
materials,  and  the  possible  potential  energy  must  be  checked  against  the  ac- 
tual energy  liberated. 

The  food  is  intended  to  supply  the  place  of  the  material  which  has  been 
utilized  by  the  body,  and,  in  a  simpler  form,  eliminated  in  the  excretions. 
But  in  the  choice  of  a  diet  this  is  not  enough;  the  food  should  be  sufficient 
to  supply  such  need  without  waste  and  without  unduly  increasing  the  output 
of  excreta,  while  at  the  same  time  the  body  should  be  maintained  in  health, 
without  increase  or  loss  of  weight.  The  food  must  also  supply  the  energy 
liberated  without  undue  waste  of  the  tissues  themselves. 

These  requisites  of  a  diet  scale  then  allow  for  wide  alterations  in  the 
amount  of  different  kinds  of  foods  under  different  circumstances.  Numer- 
ous and  most  valuable  experiments  have  been  performed  in  recent  years  to 
determine  just  what  each  article  of  the  common  food  materials  contributes  to 
the  growth  of  the  tissues  and  to  the  kinetic  energy  liberated  by  the  tissues. 
The  potential  energy  of  the  food  can  also  be  checked  against  the  kinetic 
energy  liberated.  A  single  illustration  of  this  class  will  serve.  In  an  experi- 
ment with  mixed  food  lasting  through  four  days,  on  a  man  with  body  weight 
of  64  kilograms,  and  doing  a  minimum  amount  of  work,  Atwater  made  the 
following  determinations: 

454 


METABOLISM,    NUTRITION,    AND   DIET  455 

WEIGHT,  COMPOSITION,  AND  HEAT  OF  COMBUSTION  OF  FOODS  AND  EXCRETA 

PER  DAY. 


1 

Si 

1 

c 

i 

}! 

I 

j 

Hydrogen. 

8  g 
8 

Food 

Beef 

Grms. 

Grms. 
6  1     2 

Grms. 

Grms. 
31 

Grms. 

Grms. 
S    62 

Grms. 

20  .  o  s 

Grms. 
2  .  OO 

Calor- 
ies 

227 

Butter 

2  5 

2     6 

5 

2  I  .  I 

.08 

I  c.  77 

2  .  50 

104 

Whole  milk      • 

850 

726.8 

3*   3 

45-  r 

79  .9 

5.  10 

67  .  74 

0.04 

768 

Bread 

•7  OO 

123.9 

22.2 

12  .  O 

139  .  2 

•?  .  oo 

82.53 

12  .  24 

835 

Shredded    wheat 
biscuit. 
Ginger  snaps  .... 
Sugar  

5° 
5° 

2O 

4.1 

3-3 

4.8 

2.8 

•7 
3-6 

39-7 

39-2 
20  .0 

.84 

•5° 

20  .46 

21.12 
8.42 

2.87 

3-07 
i  .  ^o 

204 
212 

79 

Total     food     per 
day. 

-.395 

921.9 

97-7 

85.6 

278.0 

16.04 

236.09 

34.82 

2,519 

Average  feces  per 
day. 
Average  urine  per 
day. 
Excretions  —  lungs 

98.8 
1420.8 

77-7 
1363.0 
881  .0 

7-7 

4.0 

6-3 

1.23 
I5-85 

9.98 
11.79 

221  .  <; 

1.42 
2.98 

no 

135 

2,397 

and  skin. 

Total  excreta  per 
day. 



2322  .6 

17.08 

243-27 

4.40 

2,642 

Balance 

—i  .  04 

—7.18 

+  30.42 

—  123 

Careful  analyses  of  the  excreta,  many  of  which  we  have  already  had  oc- 
casion to  call  attention  to,  show  that  they  are  made  up,  besides  water,  chiefly 
of  the  chemical  elements  carbon,  hydrogen,  oxygen,  and  nitrogen,  but  that 
they  also  contain,  to  a  less  extent,  sulphur,  phosphorus,  chlorine,  potassium, 
sodium,  calcium,  magnesium,  iron,  and  certain  other  of  the  elements.  Since 
this  is  the  case  it  must  be  evident  that,  to  balance  this  waste,  foods  must  be 
supplied  containing  all  these  elements  to  a  certain  degree,  and  some  of 
them  in  large  amount,  viz.,  those  which  take  a  principal  part  in  forming 
the  excreta. 

The  waste  products  of  the  body  are  eliminated  through  the  lungs,  the 
skin,  the  alimentary  canal,  and  the  kidneys.  In  the  lungs  the  chief  waste 
product  is  water  and  carbon  dioxide  gas.  Some  carbon  dioxide  gas  and  small 
quantities  of  urea  and  salts  are  eliminated  through  the  skin.  From  the 
alimentary  canal  there  are  lost,  through  the  feces,  the  indigestible  and  un- 


456 


METABOLISM,    NUTRITION,    AND    DIET 


absorbed  substances  from  the  food,  together  with  products  secreted  into  the 
canal  by  the  liver,  pancreas,  and  mucous  membrane.  The  secretion  lost 
daily  by  the  kidney,  aside  from  a  large  quantity  of  water,  consists  of  nitrog- 
enous waste  products,  chiefly  urea,  and  inorganic  solids,  as  were  mentioned 
in  the  chapter  on  Excretion. 

The  relations  between  the  amounts  of  the  chief  elements  contained  in  these 
various  excreta  in  twenty-four  hours  may  be  thus  summarized: 


Water. 

c. 

H. 

N. 

O. 

By  the  lungs 

7  •JQ 

248   8 

By  the  skin  

660 

2.6 

"o1  •  l  5 

72 

By  the  urine.  .... 

I  708 

Q   8 

3*) 

i  <  8 

.  * 

By  the  feces 

I  20 

20   o 

•  o 

A  0  •  ° 

•  u 

•  u 

Grams      .  . 

2  818 

28l     2 

67 

18   8 

681    A  c 

From  the  water  in  this  table  should  be  subtracted  the  296  grams  of  water 
which  are  produced  by  the  union  of  hydrogen  and  oxygen  in  the  body  during 
the  process  of  oxidation,  and  there  should  be  added  to  the  respective  columns 
the  corresponding  amounts  of  the  constituent  elements,  i.e.,  33  grams  of 
hydrogen  and  262  grams  of  oxygen.  There  are  26  grams  of  salts  eliminated 
through  the  urine,  and  6  by  the  feces;  a  total  of  32  grams. 

The  quantity  of  carbon  daily  lost  from  the  body  amounts  to  about  281 . 2 
grams  and  of  nitrogen  18.8  grams,  and  if  a  man  could  be  fed  by  these  ele- 
ments, as  such,  the  problem  would  be  a  very  simple  one;  a  corresponding 
weight  of  charcoal  and,  allowing  for  the  oxygen  in  it,  of  atmospheric  air 
would  be  all  that  is  necessary.  But  an  animal  can  live  upon  these  elements 
only  when  they  are  arranged  in  a  particular  manner  with  others,  in  the  form 
of  such  food  stuffs  as  we  have  already  enumerated,  page  322  et  seq.;  more- 
over, the  relative  proportion  of  carbon  to  nitrogen  in  either  of  these  com- 
pounds alone  is  by  no  means  the  proportion  required  in  the  diet  of  man. 
Thus,  in  protein,  the  proportion  of  carbon  to  nitrogen  is  only  as  3.5  to  i. 
If,  therefore,  a  man  took  into  his  body,  as  food,  sufficient  protein  to  supply 
him  with  the  needed  amount  of  carbon,  he  would  receive  more  than  four  times 
as  much  nitrogen  as  is  needed;  and  if  he  took  only  sufficient  to  supply  him 
with  nitrogen,  he  would  be  starved  for  want  of  carbon.  It  is  plain,  therefore, 
that  he  should  take  with  the  albuminous  part  of  his  food,  which  contains  so 
large  an  amount  of  nitrogen  in  proportion  to  the  carbon  he  needs,  substances 
in  which  the  nitrogen  exists  in  relatively  much  smaller  quantities  than  the 
carbon. 


METABOLISM    OF    PROTEINS 


457 


It  is,  therefore,  evident  that  the  diet  must  consist  of  several  compounds, 
not  of  one  alone. 

Many  valuable  observations  have  been  made  with  a  view  of  ascertaining 
the  effect  upon  the  metabolism  of  a  variation  in  the  amount  and  nature  of 
food.  These  are  of  great  assistance  in  the  consideration  of  dietetics. 


METABOLISM  OF  PROTEINS. 

Nitrogenous  Equilibrium. — Experiments  have  been  made,  to  a  con- 
siderable extent  upon  dogs,  which  demonstrate  the  necessity  for  protein 
food.  After  a  preliminary  period  without  food,  during  which  the  output  of 
nitrogen  as  shown  by  the  urea  has  diminished  to  a  comparatively  constant 
amount,  an  animal  is  fed  with  a  diet  of  lean  meat  which  would  suffice  to  pro- 
duce the  amount  of  urea,  and  so  of  flesh,  which  it  has  been  losing  during  its 
starvation  period.  The  effect  of  this,  however,  is  at  once  to  send  up  the 
amount  of  urea  excreted  to  a  point  above  that  which  had  been  lost  previous 
to  the  commencement  of  the  flesh  diet.  Thus  the  output  of  nitrogen  still 
exceeds  its  income,  and  the  weight  of  the  animal  continues  slowly  to  dimin- 
ish. It  is  only  after  a  considerable  increase  of  the  flesh  given  in  the  food  that 
a  point  is  reached  where  the  income  and  the  expenditure  of  nitrogen  are  equal, 
and  at  which  the  animal  is  not  using  up  or  storing  the  nitrogen  in  its  own 
tissue,  and  is  no  longer  losing  flesh.  This  condition  in  which  the  nitrogen 
of  the  egesta  equals  the  nitrogen  of  the  ingesta  is  known  as  nitrogenous 
equilibrium. 

EXPERIMENT  IN  NITROGENOUS  EQUILIBRIUM. 


Days  of  experiment. 

N 
Intake. 
Grams. 

N 
Output. 
Grams. 

Per  cent. 
Differ- 
ence. 

S 
Intake. 
Grams. 

S 

Output. 
Grams. 

Tic 

QO     OO 

80  .  81 

—  O  .  2  I 

*••      L      3  
A      T2 

III     60 

I  •?  2     7  ^ 

+  o  .88 

UT       2 

•?  c   80 

^6    16 

+  1  .00 

O  0  '  ou 

•TA-2        T  7 

—  0.86 

3    JI  

ml     7 

I  ZA    8  I 

I  C3,  .  02 

—  o.  si 

8-17  

213.72 

213.26 

—  O  .  21 

12.77 

12.79 

In  the  dog,  according  to  Waller,  nitrogenous  equilibrium  does  not  occur 
until  the  amount  of  flesh  of  the  food  is  over  three  times  as  great  as  would  be 
necessary  to  supply  the  nitrogen  of  the  urine  during  a  period  of  starvation. 
Thus  a  dog  excretes  during  a  starvation  period  0.50  gram  of  urea  per  kilo 


458  METABOLISM,    NUTRITION,    AND    DIET 

of  body  weight.  In  order  to  satisfy  this  waste  it  would  be  necessary  to 
administer  1.50  grams  per  kilo  of  meat  protein.  This  at  once  increases  the 
urea  excreted  to  about  0.75  gram  per  kilo  of  body  weight,  and  nitrogenous 
equilibrium  is  not  attained  until  over  three  times,  viz.,  3  grams  per  kilo 
of  body  weight,  of  meat  protein  is  given.  Foster  gives  even  a  larger 
figure.  The  immediate  effect,  therefore,  of  increasing  the  protein  food  is 
largely  to  increase  the  excretion  of  urea. 

Studies  in  nitrogenous  equilibrium  are  based  on  the  fact  that  when  an 
animal  is  given  a  diet  with  a  constantly  increasing  amount  of  protein  food 
from  day  to  day,  after  a  few  days  the  total  nitrogen  found  in  the  excreta 
exactly  balances  that  taken  in  the  food.  This  condition  of  nitrogenous 
equilibrium  is  established  at  different  levels,  varying  sometimes  according 
to  the  individual  and  with  the  kind  and  quantity  of  other  food  principles 
taken  at  the  same  time  as  the  nitrogenous  foods. 

Chittenden's  later  metabolism  experiments  have  shown  that  with  free 
choice,  but  moderate  use,  of  accessory  articles  of  diet,  the  human  body  can 
maintain  itself  in  nitrogenous  equilibrium  for  at  least  several  months  on  an 
average  of  6  to  10  grams  of  nitrogen  per  day,  the  equivalent  of  37.5  to  62.5 
grams  of  dry  protein  or  four  times  as  much  lean  beef. 

The  Role  of  Proteins  in  Metabolism. — The  proteins  of  food  are 
described  by  Voit  as  having  two  relations  to  the  protein  metabolism,  also  to 
outgoing  urea.  The  first  part  of  the  protein  of  the  food  goes  to  maintain  the 
ordinary  and  quiet  metabolism  of  the  tissues,  for  which  purpose  it  is  actually 
built  up  into  the  living  protoplasmic  molecule.  The  second  part  being 
more  directly  oxidized  causes  a  more  rapid  formation  of  urea,  but  never 
becomes  a  part  of  the  actual  protoplasmic  molecule.  The  former  proteins 
are  called  mor photic  or  tissue  proteins,  the  latter  circulating  or  floating  pro- 
teins. Normally  more  protein  is  eaten  than  is  needed  to  supply  protein 
to  the  protoplasm  for  growth,  as  has  just  been  stated.  Pfliiger  takes  the 
view,  however,  that  the  tissues  must  have  an  excess  of  protein  to  destroy  in 
order  to  perform  their  metabolic  processes  normally.  This  use  of  the 
proteins  to  form  heat  by  their  oxidation,  and  not  to  produce  tissue,  was 
looked  upon  by  the  older  physiologists  as  a  wasteful  use  of  good  material, 
and  was  called  luxus  consumption.  This  use  is  now  disproved. 

Folin  has  recently  announced  a  theory  of  protein  metabolism  in  which 
he  calls  special  attention  to  the  relation  of  the  nitrogenous  excretion  products 
to  the  nitrogenous  intake.  He  has  presented  evidence  to  show  that  the  urea 
contained  in  the  urine  varies  almost  directly  with  the  quantity  of  protein  in 
the  food;  that  the  ammonia  also  varies  with  the  protein  in  the  food;  that  the 
uric  acid  decreases  (and  increases)  with  the  protein  in  the  food,  but  not  in 
direct  ratio;  while  the  creatinin  excreted  is  " wholly  independent  of  quan- 
titative changes  in  the  total  amount  of  nitrogen  eliminated." 


PROTEINS    AS    FAT-    AND    AS    GLYCOGEN-FORMERS  459 

TABLE  SHOWING  THE  OUTPUT  OF  NITROGEN  IN  A  NORMAL,  HEALTHY  INDIVID- 
UAL ON  A  FOOD  RICH  IN  NITROGEN,  JULY  13™,  AND  POOR  IN  NITROGEN 
JULY  20TH  (FOLIN). 

July   i3th.  July  2oth. 

Volume  of  urine  .  .  .1,170  c.c. .  385  c.c. 

Total  nitrogen 1 6 . 08  grams  3  .  60  grams 

Urea  nitrogen 14.  70  grams  =87  .  5  per  cent.  22.0    grams  =61 .  7  per  cent. 

Ammonia  nitrogen .    0.49  grams  =    3.0  per  cent.  0.42  grams  =  n  .3  per  cent. 

Uric-acid    nitrogen.    o.i8grams=    i .  i  per  cent.  0.09  grams  =    2  .  5  per  cent. 

Creatinin   nitrogen .    0.58  grams  =    3.6  per  cent,  o .  60  grams  =  17.2  per  cent. 

Undetermined  nitro-  0.85  grams  =   4.9  per  cent.  0.27  grams  =    7.3  per  cent, 
gen. 

Folin  states  this  theory  as  follows:  "It  is  clear  that  the  metabolic  proc- 
esses resulting  in  the  end  products  which  tend  to  be  constant  in  quantity 
appear  to  be  indispensable  for  the  continuation  of  life;  or,  to  be  more  defi- 
nite, those  metabolic  processes  probably  constitute  an  essential  part  of  the 
activity  which  distinguishes  living  cells  from  dead  ones.  I  would  therefore 
call  the  protein  metabolism  which  tends  to  be  constant,  tissue  metabolism, 
or  endogenous  metabolism;  the  other,  the  variable  protein  metabolism,  I 
would  call  the  exogenous  or  intermediate  metabolism. 

"The  endogenous  metabolism  sets  a  limit  to  the  lowest  level  of  nitrogen 
equilibrium  attainable.  Just  where  that  level  is  fixed  will  depend  on  how 
much,  if  any,  urea  is  derived  from  the  same  katabolic  processes  that  produce 
the  creatinin.  If  this  can  be  determined,  we  shall  have  a  formula  expressing 
more  or  less  definitely  the  point  of  lowest  attainable  protein  katabolism, 
because  at  such  a  point  the  percentage  composition  of  the  urine  should  be 
•practically  constant.  The  total  nitrogen  eliminated  when  this  constant  com- 
position of  the  urine  has  been  reached  will  indicate  the  lowest  attainable 
level  of  nitrogen  equilibrium. " 

The  condition  of  nitrogenous  equilibrium,  therefore,  is  one  which  may 
be  maintained  even  if  the  amount  of  protein  taken  as  diet  far  exceeds  the 
necessities  of  the  economy,  the  urea  being  excreted  in  excessive  amount. 
The  wasteful  use  of  protein  food  which  is  so  prevalent  may  not  be  attended 
with  harmful  consequences,  so  long  as  the  excreting  organs  are  able  to 
eliminate  nitrogen  from  the  body  but  it  overworks  those  organs. 

It  is  only  in  cases  of  growth,  by  putting  on  of  flesh,  as  in  growing  children, 
that  nitrogen  is  retained  in  the  body  in  health,  except  to  a  very  small  amount. 
According  to  calculations  which  have  been  made,  it  appears  that  the  body 
puts  on  30  grams  of  flesh  for  every  gram  of  nitrogen  so  retained. 

Proteins  as  Fat-  and  as  Glycogen-Formers. — Protein  food  is  un- 
doubtedly a  source  of  energy  in  the  body;  and  one  can  say  that  such  protein 
as  is,  according  to  Voit's  view,  metabolized  without  becoming  part  of  the 
tissue  may  be  considered  a  source  of  energy.  If  this  be  true,  one  might  ex- 


460  METABOLISM,    NUTRITION,    AND    DIET 

pect  that  proteins  could  be  metabolized  into  other  forms,  such  as  carbo- 
hydrates and  fats.  Bernard  long  ago  stated  that  protein  was  a  glycogen- 
former;  that  abundant  glycogen  was  stored  in  the  liver  when  flesh  diet  was 
fed,  and  argued  that  protein  was  the  source  of  the  glycogen.  The  careful 
work  of  a  number  of  investigators  has  not  obtained  sufficient  evidence  to 
clear  up  this  question  absolutely,  but  the  weight  of  evidence  is  in  favor  of  the 
view  that  in  the  body  sugar  can  be  formed  from  proteins.  Lusk  has  recently 
shown  that  some  of  the  amino  acids  must  have  been  converted  into  dextrose 
after  deamidization  in  the  body.  Whether  or  not  protein  can  be  metabolized 
into  fat,  and  stored  as  such,  seems  at  present  an  open  question,  notwith- 
standing the  immense  amount  of  work  expended  in  trying  to  solve  the 
problem. 

Cramer  fed  450  grams  of  lean  meat  per  day  to  a  cat  in  a  respiration 
chamber  for  8  days.  The  daily  excretion  of  nitrogen  was  13  grams,  of 
carbon  34.3  grams;  calculating  the  amount  of  carbon  in  the  food  as  41.6 
grams  daily,  this  would  leave  7 . 3  grams  retained.  This  carbon  might  be 
stored  in  the  form  of  glycogen  or  as  fat.  Calculated  as  glycogen,  it  gives  an 
amount  greater  than  an  animal  of  that  size  could  retain.  Therefore,  the 
probabilities  are  that  the  carbon  is  deposited  in  the  form  of  fat. 

In  the  examination  of  the  fat  formed  in  the  larvae  of  blow-flies  developing 
in  a  quantity  of  coagulated  blood,  Hoffmann  found  ten  times  more  fat  than 
existed  in  the  blood.  These  experiments  point  in  the  direction  of  fat 
formation  from  protein. 

The  Effect  of  a  Gelatin  Diet. — The  albuminoid  eaten  in  greatest 
quantity  is  gelatin.  Though  gelatin  closely  resembles  the  protein  mole- 
cule chemically,  it  cannot  replace  entirely  the  protein  of  the  food.  As  was 
stated  in  the  chapter  on  the  Chemistry  of  the  Body,  gelatin  is  deficient  in 
certain  amino  acids,  notably  tryptophane  which  is  not  present  at  all. 
It  is  probable  that  because  of  the  absence  of  this  "building  stone,"  the 
body  tissues  cannot  reform  their  characteristic  proteins  for  which  this 
amino  acid  would  be  an  essential  constituent.  In  other  words,  nitrogenous 
equilibrium  cannot  be  maintained  on  a  diet  consisting  of  gelatin,  carbo- 
hydrates, and  fats.  Gelatin,  then,  is  a  substance  whose  food  value  in  part 
is  comparable  to  that  of  carbohydrates  and  fats,  as  the  following  experiments 
will  prove:  On  a  diet  of  500  grams  of  meat,  without  any  gelatin,  the  subject 
lost  nitrogen  to  the  equivalent  of  22  grams  of  protein,  but  when  200  grams 
of  gelatin  were  added  the  subject  gained  54  grams.  In  another  experiment, 
when  the  diet  consisted  of  2,000  grams  of  meat  without  gelatin,  the  gain 
was  the  equivalent  of  30  grams  of  protein,  but  when  200  grams  of  gelatin 
were  added  the  gain  became  376  grams.  The  lack  of  a  mixed  protein  food 
value  is  proven  by  still  a  third  experiment  in  which  the  diet  consisted  at 
first  of  200  grams  each  of  meat  and  of  gelatin;  here  the  gain  was  the  equiva- 
lent of  25  grams  of  protein,  but,  when  the  meat  was  omitted  and  the  gelatin 


THE    FORMATION    OF    UREA  461 

alone  given,  there  was  a  loss  of  118  grams.  In  these  cases  gelatin  did  not 
take  the  place  of  protein  in  any  sense,  buit  rather  saved  it  from  oxidation  as 
a  source  of  energy.  The  protein  was  so  protected  that,  instead  of  being  used 
up,  it  helped  to  form  tissue  and  increased  the  body  weight.  Gelatin,  there- 
fore, saved  the  protein  material  for  constructive  processes. 

Murlin  has  investigated  more  exactly  the  substitution  of  gelatin  for  the 
mixed  proteins  in  the  food.  In  a  series  of  experiments  on  dogs,  the  nitrogen 
output  was  first  determined  during  fasting  periods.  Varying  amounts 
of  gelatin  containing  from  a  fourth  to  two-thirds  of  this  amount  of  nitrogen 
were  then  fed,  the  remaining  three-fourths  to  one-third  of  the  fasting  quantity 
being  supplied  in  meat  or  other  proteins.  The  calorific  requirement  of 
the  animal  was  made  up  in  each  experiment  with  fats  and  carbohydrates. 
Results  show  an  equal  sparing  of  body  protein,  whether  one-fourth,  one- 
third,  or  one-half  of  the  fasting  nitrogen  was  fed  in  the  form  of  gelatin, 
the  coincident  sparing  of  protein  by  fats  and  carbohydrates  being  the  same. 
When  the  coincident  sparing  of  protein  by  non- nitrogenous  food  was  in- 
creased by  feeding  a  larger  percentage  of  carbohydrates  and  less  fat,  the 
fraction  of  the  fasting  nitrogen  fed  in  the  form  of  gelatin  could  be  raised  to 
two-thirds,  the  other  one-third  being  fed  in  meat.  Nitrogenous  equilibrium 
was  maintained  on  this  diet  for  several  days.  The  same  result  was  obtained 
on  man.  The  evidence  at  hand  indicates  that  other  individual  proteins, 
in  which  certain  amino  acid  constituents  are  deficient  in  quantity  or  which 
are  absent,  like  gelatin  would  not  replace  entirely  the  ordinary  mixed 
protein  requirements. 

The  Formation  of  Urea. — The  nitrogenous  fraction  of  the  protein  mole- 
cule is  in  the  end  converted  largely  into  urea  and  is  excreted  from  the  body  in 
that  form,  as  described  in  the  chapter  on  Excretion.  The  method  of  forma- 
tion of  urea  as  well  as  the  place  where  this  occurs  has  given  rise  to  great 
controversy,  while  the  intermediate  products  between  proteins  and  urea 
have  not  as  yet  been  fully  determined.  We  can  state  with  certainty  that 
urea  is  not  formed  in  the  kidneys,  since  it  is  not  only  found  in  the  blood  of 
the  renal  artery,  but  it  accumulates  in  the  blood  if  the  kidneys  are  diseased 
or  removed  and  the  separation  of  the  urine  is  interfered  with.  Circulation 
of  blood  through  the  kidney  does  not  result  in  the  formation  of  more  urea 
than  is  present  in  the  blood  to  begin  with. 

There  are  a  number  of  experiments  that  prove  that  urea  is  formed  in  the 
liver.  The  power  of  the  liver  cells  to  form  urea  is  shown  by  the  increase  of 
urea  in  the  blood  leaving  an  isolated  living  liver  through  which  an  artificial 
circulation  is  kept  up.  When  ammonium  carbamate  and  other  ammonium 
salts  are  added  to  the  blood,  the  urea  increases  more  rapidly  and  to  a  greater 
extent.  This  change  occurs  even  when  the  living  hepatic  tissue  is  chopped 
up  and  simply  mixed  with  the  ammonium  compounds  in  a  beaker. 

If  blood  from  a  well-fed  animal  be  circulated  through  the  isolated  liver, 


462  METABOLISM,    NUTRITION,    AND    DIET 

there  is  a  distinct  increase  in  the  amount  of  urea  it  contains.  On  the  other 
hand,  if  the  blood  be  from  a  fasting  animal  there  is  little  or  no  increase  of 
urea.  Evidently,  then,  the  blood  from  a  well-fed  animal  contains  something 
which  the  liver  cells  are  capable  of  transporting  into  urea.  And,  finally,  if 
the  liver  be  removed  and  the  animal  kept  alive,  as  has  been  done  by  Pawlow, 
there  is  a  marked  diminution  in  the  quantity  of  urea  in  the  urine.  The 
power  of  the  liver  to  form  urea  is  thus  demonstrated.  The  question  which 
now  presents  itself  is,  what  is  this  antecedent  substance  or  substances? 

It  has  already  been  indicated  that  urea  follows  closely  the  amount  of 
protein  taken  with  the  food,  hence  we  must  look  directly  to  the  nitrogenous 
fraction  of  protein  cleavage  as  the  final  source  of  urea.  While  the  different 
steps  in  the  process  of  cleavage,  probably  hydrolytic  (Folin),  are  yet  very 
obscure,  still  it  is  believed  that  ammonia  is  split  off  from  the  protein  cleav- 
age products  and  is  then  built  up  into  urea  by  the  liver.  It  is  now  believed 
that  ammonium  carbamate  is  at  least  one  true  antecedent  of  urea. 

In  these  experiments  the  liver  is  first  shut  out  of  the  general  circulation 
by  an  Eck's  fistula  connecting  the  portal  vein  with  the  vena  cava.  This 
operation  cuts  off  the  chief  blood  supply  of  the  liver,  viz.,  the  portal  blood, 
but  it  leaves  the  small  hepatic  artery  with  its  oxygen  supply  to  the  liver. 
When  animals  survive  this  operation  it  is  found  that  they  can  live  only  when 
fed  very  carefully  on  a  mixed  diet  from  which  proteins  are  almost  entirely 
eliminated,  and  that,  if  the  food  contain  an  excess  of  proteids,  convulsions 
ensue  with  fatal  termination.  Investigation  of  the  composition  of  the  urine 
and  of  the  blood,  with  the  Eck's  fistula,  shows  that  the  end  product  of  pro- 
tein metabolism  is  represented  by  ammonium  carbonate  and  carbamate  and 
that  there  is  a  considerable  decrease  in  urea.  If  ammonium  salts  are  in- 
jected into  the  blood  of  normal  animals  in  a  larger  quantity  than  the  liver 
can  dispose  of,  death  ensues,  following  convulsions  of  the  same  nature  as 
those  produced  by  an  excess  of  protein  food  in  the  animals  operated  on. 

Ammonium  carbamate  is  shown  to  be,  in  part  at  least,  the  direct  ante- 
cedent of  urea.  The  reaction  by  which  the  liver  changes  it  to  the  inert 
form  of  urea  is  as  follows: 

O.NH4  NH2  NH2 

O:C/         —  H2O    —    O:C<^         -H2O  —  O:C/ 

O.NH4  O.NH4  NH2 

Ammonium  Ammonium  Urea 

carbonate  carbamate 

The  elimination  of  urea  is  increased  very  slightly  by  muscular  activity. 
But  there  is  no  direct  relationship  between  the  amount  of  work  done  and  the 
amount  of  nitrogen  excreted  as  urea. 

There  is  experimental  evidence  to  show  that  while  the  liver  produces  the 


THE    METABOLISM    OF    FATS  463 

major  part  of  the  urea  eliminated,  other  organs  or  tissues  are  capable  of 
forming  it  to  a  limited  degree. 

Formation  of  Uric  Acid. — The  relation  which  uric  acid  and  urea  bear 
to  each  other  in  different  animals,  as  we  have  seen,  is  still  obscure.  The 
fact  that  they  exist  together  in  the  same  urine  makes  it  seem  probable  that 
they  have  different  origins.  The  entire  replacement  of  one  by  the  other,  as 
of  urea  by  uric  acid  in  the  urine  of  birds,  serpents,  and  many  insects,  shows 
their  close  relationship.  But  although  it  is  true  that  uric  acid  on  oxidation 
yields  urea,  this  is  not  evidence  that  uric  acid  is  an  antecedent  of  urea  in  the 
nitrogenous  metabolism  of  the  body.  The  chemical  structure  of  the  uric 
acid  shows  it  has  a  nucleus  of  purin,  and  therefore  is  a  close  relative  of 
adenin,  guanin,  hypoxanthin,  xanthin,  theobromin,  caffein,  etc.  The  nu- 
cleins  on  cleavage  yield  members  of  this  group,  hence  may  be  looked  to  as 
the  primary  source  of  uric  acid  in  man.  Uric  acid,  according  to  Chittenden, 
has  a  double  origin — endogenous  from  nuclear  metabolism,  and  exogenous 
from  metabolism  of  foods  rich  in  nuclear  and  other  purin  compounds.  In 
man,  at  least,  the  uric  acid  is  to  be  ascribed  to  these  two  sources. 

Operative  experiments  on  birds  tend  to  show  that  the  final  step  in  uric- 
acid  formation  takes  place  chiefly  in  the  liver,  for  on  the  removal  of  this  organ 
ammonium  compounds,  i.e.,  lactates,  accumulate  in  the  blood. 

Hippuric  Acid,  Creatinin. — The  hippuric  acid  found  in  the  urine 
is  derived  in  part  from  aromatic  constituents  of  vegetable  diet  which  can  be 
transformed  into  benzoic  acid  in  the  body.  It  is  derived  in  part  also  from 
the  phenyl  propionic  acid  formed  in  the  intestinal  putrefaction  of  protein. 
Hippuric  acid  is  formed  from  the  union  of  benzoic  acid  with  glycocoll 
(C2H5NO2  +  C7H6O2  =  C9O9NO3  +  H2O).  This  union,  in  dogs,  takes 
place  under  experimental  conditions  in  the  kidneys  themselves,  but  in  other 
animals  the  synthesis  will  occur  after  nephrectomy. 

The  source  of  the  nitrogenous  extractives  of  the  urine  is  chiefly  from  the 
metabolism  of  the  nitrogenous  foods  and  tissues,  but  we  are  unable  to  say 
whether  these  nitrogenous  bodies  have  merely  resisted  further  decomposition 
into  urea,  or  whether  they  are  the  representatives  of  the  decomposition  of 
special  tissues,  or  of  special  forms  of  metabolism  of  the  tissues.  There  is, 
however,  one  exception,  and  that  is  in  the  case  of  creatinin.  This  represents 
not  only  the  creatinin  which  enters  the  body  in  ordinary  flesh  food,  but  is  a 
nitrogenous  waste  which  Folin  regards  as  a  measure  of  muscle  metabolism. 
The  creatinin  eliminated  is  almost  a  constant  quantity  in  a  given  individual, 
irrespective  of  the  quantity  of  protein  in  the  diet.  Koch  has  shown  some 
relation  of  creatinin  excretion  to  the  amount  of  lecithin  in  the  food. 

THE  METABOLISM  OF  FATS. 

Fats,  with  carbohydrates,  are  the  direct  source  of  most  of  the  energy 
manifested  by  the  body,  a  fact  demonstrated  by  numerous  observations. 


464 


METABOLISM,    NUTRITION,    AND    DIET 


The  Energy  Value  of  Fats  in  Metabolism.— Fats,  in  comparison 
with  other  food  principles,  are  of  especial  value  as  sources  of  energy.  They 
are  completely  oxidized  in  the  body  to  carbon  dioxide  and  water,  and  yield, 
therefore,  as  much  energy  to  the  body  as  they  yield  upon  oxidation  outside 
the  body.  The  energy  equivalent  of  i  gram  of  fat  is  9 . 3  large  Calories, 
more  than  twice  that  of  starch,  which  in  the  body  yields  only  4.1  Calories 
per  gram,  or  of  protein  with  4.1  Calories  available  yield  of  energy. 

A  study  of  the  elimination  of  nitrogen  and  of  carbon  during  fasting  shows 
that  the  fats  contribute  to  energy  formation  for  many  days.  This  is  illus- 
trated by  the  following  computation  by  Voit: 

METABOLISM  IN  A  DOG  DURING  FASTING.     (VoiT.) 


Loss  per  kilogram  of  live  weight. 

Proteins 
in  grams 

Fats 
in  grams 

Total 
weight 

Second  day 

2.21 

2  .  62 

7.2  .87 

Fifth  day 

i    1  3 

?  .  2  S 

•2  I       67 

Eighth  day               .  .                        .  .           ... 

o  .  06 

T>  '  2  ^ 

•?O  .  <?4 

The  amount  of  fat  metabolized  is  sharply  influenced  by  the  amount  and 
kind  of  other  food.  For  example,  if  the  amount  of  fat  metabolized  per  day 
in  fasting  is  first  determined,  then  a  ration  of  protein  given  for  a  few  days, 
followed  by  a  second  fasting  period,  it  will  be  found  that  the  metabolism  of 
body  fat  is  sharply  increased  in  the  second  period,  due  to  the  stimulating 
influence  of  the  protein.  This  is  demonstrated  by  the  following  determi- 
nation of  Rubner: 


Food  of  dog. 
(Rubner.) 

Nitrogen  of 
food. 

Nitrogen 
excreted. 

Body  fat 
metabolized. 

o  . 

o  . 

4.38 

4Q  •  33 

41  c 

grams  lean  meat  . 

14.11 

17.72 

2  c  .  44  average. 

o 

o 

2  .  80 

70  .  04 

760 

grams  lean  meat  

2  5  .  l6 

2O  .  6^ 

30.73  first  two  days. 

The  fat  of  the  ordinary  daily  diet  is  absorbed  into  the  blood  and  no  doubt 
contributes  directly  to  oxidation  processes.  In  some  of  the  lower  carnivor- 
ous vertebrates  fats  are  undoubtedly  the  primary  and  direct  source  of  the 
energy  of  daily  activity.  Just  the  steps  in  this  oxidation  process  cannot  at 
present  be  given.  If  the  fat  of  the  food  is  insufficient,  then  the  body  store 
is  immediately  drawn  upon;  if  in  excess,  then  it  is  stored  in  the  body.  This 


SOURCE    OF    THE    BODY    FAT 


465 


observation  is  strongly  supported  by  the  work  of  Eckles  on  the  variation  in 
the  percentage  of  fat  in  the  milk  of  dairy  cows  under  the  influence  of  tempo- 
rary short  feed.  If  a  cow  is  given  an  insufficient  amount  of  food  immediately 
the  percentage  of  fat  in  the  milk  sharply  increases,  sometimes  amounting  to 
almost  double  the  original  fat  content  of  the  milk. 

Source  of  the  Body  Fat. — Excess  of  fat  in  the  food  can  be  stored  as 
fat  in  the  body.  This  fact  is  demonstrated  by  Voit,  Hoffmann,  Rubner, 
and  others.  Rubner  states  that  82  to  92  per  cent,  of  the  fat  excess  can  be 
stored.  The  fat  stored  was  long  thought  to  be  the  same  kind  given  in  the 
food,  even  though  the  usual  fat  of  the  animal  was  different.  The  melting 
point  of  dog's  fat  is  about  20°  C,  but  by  feeding  an  excess  of  mutton  fat  the 
melting  point  has  been  raised  to  40°  C.  The  subcutaneous  fat  of  pigs  sub- 
jected to  this  experiment  is  more  or  less  fluid  according  to  the  melting  point 
of  the  fat  fed.  However,  we  now  know  that  this  conception  is  only  partially 
correct.  The  physical  and  chemical  constants  show  that  the  fat  laid  down 
approaches  in  characteristics  the  fat  fed,  but  that  it  is  not  the  same.  Even 
in  the  alimentary  canal  during  digestion,  changes  occur  which  modify  the 
melting  point  and  other  properties  of  remnants  of  the  fat  not  absorbed.  Fat 
that  is  resynthesized  in  the  mucous  epithelium  after  absorption  is  similar 
but  not  identical  with  the  fats  of  the  food.  This  possibly  rests  upon  a  varia- 
tion in  the  number  and  character  of  the  unsaturated  bonds  during  fat 
mobilization  in  the  body. 

The  body  fat  can  also  be  derived  from  carbohydrate  food,  a  fact  which 
the  practices  of  the  stock  feeder  and  dairyman  constantly  verify.  The  ex- 
periments will  present  the  matter  more  vividly  than  pages  of  description. 


GAIN  IN  FAT  OF  A  PIG  FED  ON  RICE.      (MEISSL  AND  STROHMER.) 


Pig  weight. 

Rice  fed 
daily. 

Fat  in 
food. 

Protein 
in  food. 

Protein 
gain. 

Carbon 
gain. 

Net  gain 
in  carbon. 

140    kgm. 

2  kgm. 

5  •  3  gm- 

104  gm. 

38  gm. 

289  gm. 

269   gm. 

It  is  obvious  that  the  5.3  grams  of  fat  and  the  66  grams  of  protein  cannot 
account  for  the  carbon  retained,  and  one  must  look  to  the  carbohydrate  as 
the  source  of  the  fat. 

Jordan  placed  a  Jersey  cow  on  a  feed  of  hay  and  grain  from  which  the 
fat  was  extracted.  The  cow  in  95  days  assimilated  5 . 7  pounds  of  fat,  in- 
creased 47  pounds  in  weight,  and  produced  62.9  pounds  of  fat  in  the  milk. 
The  nitrogen  excreted  was  the  equivalent  of  33.3  pounds  of  protein.  The 
non-nitrogenous  moiety  of  the  protein,  if  its  carbon  had  all  gone  into  fat, 


466  METABOLISM,    NUTRITION,    AND    DIET 

could  not  have  produced  over  17  pounds.  Summarized,  this  experiment 
shows  conclusively  that  fat  is  synthesized  from  carbohydrate.  It  requires 
about  2 . 7  grams  of  dextrose  to  form  i  gram  of  fat,  and  this  condensation 
takes  place  with  the  formation  of  carbon  dioxide  and  water  and  the  libera- 
tion of  about  1 5  per  cent,  of  the  available  heat  of  oxidation. 

Persistent  excess  of  carbohydrate  food  produces  an  accumulation  of  fat, 
which  may  not  only  be  an  inconvenience  causing  obesity,  but  may  interfere 
with  the  proper  nutrition  of  muscles,  produce  a  feebleness  of  the  action  of 
the  heart,  and  other  troubles. 

The  formation  of  fat  from  protein  is  discussed  on  page  459. 

Fat  Mobilization  in  the  Body. — Increasing  numbers  of  recent  studies  are 
throwing  new  light  on  the  phenomenon  of  mobilization  of  fat  in  the  body. 
In  1900  Kastle  and  Loevenhart  discovered  the  reversible  action  of  lipase 
which  opened  the  door  to  a  more  fitting  conception  of  how  the  body  is  enabled 
to  handle  this  ordinarily  insoluble  constituent.  Just  as  fats  must  be  dis- 
sociated in  order  to  be  absorbed,  a  step  that  converts  the  insoluble  fat  to  a 
soluble  form,  so  the  body  transference  from  one  tissue  to  another,  from 
the  storehouse  of  fats  to  the  lymph  and  blood,  or  vice  versa,  requires  that  the 
fats  must  be  in  a  soluble  form,  i.e.,  dissociated.  The  facility  with  which 
fats  can  be  mobilized  depends  upon  the  presence  and  activity  of  fat  Upases. 
These  have  been  shown  to  be  present  in  practically  all  tissues  of  the  body, 
greater  in  some,  as  in  the  liver,  certain  glands,  etc.  Under  the  influence  of 
the  enzyme  a  fairly  constant  concentration  of  fats  is  normally  present  in 
the  blood  and  in  the  body  fluids,  and  is  always  available  as  a  source  of 
energy  to  the  oxidizing  tissues.  The  above  is  the  explanation  of  the 
increase  of  fat  in  the  milk  previously  referred  to.  When  no  food  is  com- 
ing, the  animal  draws  immediately  on  the  carbohydrates  which  are  always 
present  in  the  blood.  This  store  is  rapidly  exhausted.  As  tissue  hunger 
approaches,  an  increase  in  lipase  production  undoubtedly  takes  place,  and 
this  dissolves  and  sets  in  motion  storage  fat.  Under  these  conditions  the 
amount  of  fat  in  circulation  rapidly  increases  and  becomes  available  to  the 
muscles,  glands,  etc.,  as  illustrated  by  the  secretion  of  the  mammary  gland 
where  the  output  can  be  quantitatively  tested.  In  starvation  also  the 
general  distribution  of  fat  in  the  form  of  liposomes  increases  in  many  tis- 
sues, particularly  the  glands — a  fact  readily  determined  by  studies  of  the 
microscopic  fat  in  fresh  tissues.  This  phenomenon  is  not  a  fatty  degenera- 
tion as  is  sometimes  held,  it  is  only  the  redistribution  of  fat  under  the  law 
of  reversible  action  of  lipase  to  meet  the  emergency  of  tissue  starvation. 

Lipogenesis. — Noel  Patton  has  shown  a  distinct  accumulation  of  fat  in 
the  liver  of  the  frog  in  relation  to  fat  feeding.  For  this  phenomenon 
Loevenhart  has  suggested  the  name  lip  agenesis.  The  idea  is  that  the  liver 
in  particular  serves  to  a  degree  as  a  storehouse  of  fats,  comparable  to  the 
phenomenon  of  glycogenesis  which  characterizes  the  organ.  The  liver  is 


ENERGY   VALUE  467 

relatively  rich  in  lipase,  hence  is  to  that  extent  peculiarly  fitted  for  storing 
and  distributing  fats. 

Whether  or  not  there  are  other  special  tissues  involved  in  this  imme- 
diate problem  is  not  adequately  studied.  It  is  quite  possible  that  many 
of  the  adipose  tissues  are  involved.  Certain  tissues  of  the  body  are  great 
fat  storage  tissues — subcutaneous,  intermuscular,  peritoneal,  etc.  These 
tissues  readily  yield  their  fat  in  time  of  starvation  but  it  is  not  known  that 
they  are  concerned  in  the  production  of  lipase  at  this  time. 

Destination  of  Fats,  Obesity,  Diet,  etc. — It  has  already  been  empha- 
sized that  fats  are  a  direct  source  of  energy  of  oxidation.  This  is  their 
primary  function  in  the  body.  If  fats  are  taken  in  excess  in  the  food,  or 
are  produced  in  excess  from  carbohydrates  of  the  food,  then  they  accumu- 
late in  great  quantity,  especially  in  middle  and  old  age.  This  accumu- 
lation gives  rise  to  the  condition  of  obesity  which  is  a  condition  of  excessive 
fat  storage.  In  many  of  these  cases  there  is  persistent  storing  of  fat  in  the 
presence  of  a  dief  of  low  energy  value  and  when  considerable  physical 
labor  is  expended.  It  seems  that  such  persons  have  a  perverted  or  at 
least  disturbed  fat  metabolism.  The  explanation  is  very  often  found  in 
deviation  of  function  of  some  endocrine  gland.  For  example,  in  decreased 
secretion  of  the  posterior  lobe  of  the  pituitary  there  is  an  increase  in 
sugar  tolerance  associated  with  a  tendency  to  accumulate  large  quantities 
of  fat.  Defects  of  this  character  are  met  by  attempts  to  supply  the 
deficient  internal  secretion. 

Obesity  may  result  purely  from  over  eating,  in  which  case  the  person 
must  use  self  restraint  to  restrict  the  quantity  of  food  and  the  amount  of 
fat  and  of  carbohydrates  from  which  fats  are  easily  produced. 

In  many  persons  and  under  certain  conditions,  the  oxidation  of  the  fats 
is  incomplete.  This  is  indicated  by  an  increase  in  the  intermediary  pro- 
ducts, acetone,  etc.  This  phenomenon  gives  rise  to  the  condition  of  mal- 
nutrition expressed  by  the  term  acidosis  which  belongs  to  the  field  of  path- 
ology to  which  the  reader  is  referred. 

THE   METABOLISM    OF  CARBOHYDRATES. 

Energy  Value. — The  nutritive  function  of  carbohydrates  in  the  body 
is  to  serve  as  a  source  of  energy.  They  are  oxidized,  with  the  ultimate  pro- 
duction of  carbon  dioxide  and  water,  and  must  liberate  the  same  amount 
of  energy  as  when  burned  outside  the  body,  i.e.,  4.1  Calories  per  gram. 
A  given  weight  of  dextrose,  therefore,  furnishes  a  little  less  than  half  the 
energy  of  a  corresponding  weight  of  fat. 

Carbohydrates  are  strictly  energy-formers  and  may  be  regarded  as  the 
chief  immediate  source  of  the  energy  of  oxidations,  although  they  may  be 
synthesized  into  fats  and  possibly  even  contribute  to  protein  formation. 


468  METABOLISM,    NUTRITION,    AND    DIET 

Dextrose  is  a  constant  constituent  of  the  blood  to  the  extent  of  about  o.i 
to  0.15  per  cent.  When  this  percentage  is  increased  above  0.25,  the 
dextrose  is  either  stored  as  glycogen,  i.e.,  in  the  case  of  the  portal  blood 
during  the  absorption  of  a  carbohydrate  meal,  or  eliminated  by  the 
kidney,  i.e.,  as  in  pancreatic  diabetes. 

The  Formation  of  Glycogen — Glycogenesis: — The  important  fact 
that  the  liver  normally  forms  sugar,  or  a  substance  readily  convertible 
into  it,  was  discovered  by  Claude  Bernard  in  the  following  way:  He  fed  a 
dog  for  seven  days  with  food  containing  a  large  quantity  of  sugar  and  starch; 
and,  as  might  be  expected,  found  sugar  in  both  the  portal  and  hepatic  blood. 
But  when  the  dog  was  fed  with  meat  only,  to  his  surprise,  sugar  was  still 
found  in  the  blood  of  the  hepatic  veins.  Repeated  experiments  gave  in- 
variably the  same  result.  No  excess  of  sugar  was  found  in  the  portal  vein 
under  a  meat  diet,  if  care  was  taken  to  prevent  reflux  of  blood  from  the  he- 
patic venous  system.  Bernard  found  sugar  also  in  the  substance  of  the  liver. 
It  thus  seemed  to  him  certain  that  the  liver  formed  sugar  eVen  when,  from  the 
absence  of  saccharine  and  amyloid  matters  in  the  food,  none  could  be  brought 
directly  to  it  from  the  stomach  or  intestines. 

Bernard  subsequently  found  that  a  liver  removed  from  the  body,  and 
from  which  all  sugar  had  been  completely  washed  away  by  injecting  a  stream 
of  water  through  its  blood  vessels,  contained  sugar  in  abundance  after  the 
lapse  of  a  few  hours.  This  post-mortem  production  of  sugar  was  a  fact 
which  could  be  explained  only  on  the  supposition  that  the  liver  contained  a 
substance  readily  convertible  into  sugar.  This  theory  was  proved  cor- 
rect by  the  discovery  of  a  substance  in  the  liver  allied  to  starch,  termed 
glycogen. 

Bernard's  brilliant  researches  led  him  to  announce  the  theory  that  the 
carbohydrate  which  is  periodically  absorbed  in  large  amount  is  stored  in  the 
liver  only  to  be  reconverted  to  dextrose  and  discharged  back  into  the  blood 
stream  whenever  the  percentage  in  the  blood  falls  below  a  certain  level. 
He  regarded  the  liver  as  a  storehouse  which  regulated  the  blood  dextrose 
to  a  constant  level.  This  is  the  glycogenic  function  of  the  liver. 

Source  of  Glycogen. — The  greatest  amount  of  glycogen  is  produced 
by  the  liver  upon  a  diet  of  starch  or  sugar,  but  a  certain  quantity  is,  or  at 
least  may  be,  produced  upon  a  protein  diet.  The  glycogen,  when  stored  in 
the  liver  cells,  may  readily  be  demonstrated  in  sections  of  liver  containing  it 
by  its  reaction  (red  or  port- wine  color)  with  iodine,  and,  moreover,  when  the 
hardened  sections  are  so  treated  that  the  glycogen  is  dissolved  out,  the  proto- 
plasm of  the  cell  is  so  vacuolated  as  to  appear  little  more  than  a  framework. 
There  is  no  doubt  that  in  the  liver  of  a  hibernating  frog  the  amount  of  glyco- 
gen stored  up  in  the  liver  cells  is  very  considerable. 


SOURCE    OF    GLYCOGEN  469 

AVERAGE  AMOUNT  OF  GLYCOGEN  IN  THE  LIVER  OF  DOGS  UNDER  VARIOUS 

DIETS.      (PAVY.) 

Diet.  Amount  of  glycogen  in  the  liver. 

Flesh  food 7.19  per  cent. 

Flesh  food  with  sugar 14.5     per  cent. 

Vegetable  diet,  i.e.,  potatoes  with  bread  or  barley  meal.    17  . 23  per  cent. 

The  dependence  of  the  formation  of  glycogen  on  the  kind  of  food  taken 
is  also  shown  by  the  following  results,  obtained  by  the  same  experimenter: 

AVERAGE  QUANTITY  OF  GLYCOGEN  FOUND  IN  THE  LIVER  OF  RABBITS  AFTER 
FASTING,  AND  AFTER  A  DIET  OF  STARCH  AND  SUGAR  RESPECTIVELY. 

After  three  days'  fasting Practically  absent. 

After  diet  of  starch  and  grape  sugar *  5  •  4  per  cent. 

After  diet  of  cane  sugar 16.9  per  cent. 

Glycerol  injected  into  the  alimentary  canal  may  increase  the  glycogen 
of  the  liver.  Observations  indicate  that  glycogen  may  be  formed  in  the 
turtle  liver  when  perfused  with  very  dilute  formaldehyde  solutions.  The 
diet  most  favorable  to  the  production  of  a  large  amount  of  glycogen  is  a 
mixed  diet  containing  a  large  amount  of  carbohydrate,  but  with  some 
protein.  Glycogen  is  stored  in  other  organs  of  the  body.  Of  these  the 
muscles  are  deserving  of  special  mention.  The  amount  of  glycogen  in  the 
muscles  of  young  animals  is  often  considerable.  The  placenta  is  also  a 
storehouse  of  glycogen. 

Glycogenesis  Controlled  by  Hormone. — Glycogenesis  and  its  storage 
is  strictly  dependent  on  chemical  control.  In  disease  of  the  pancreas 
leading  to  its  degeneration,  or  in  surgical  removal  of  the  pancreas  the 
liver  is  unable  to  store  glycogen.  Even  when  the  blood  is  rich  in  sugar 
to  the  level  at  which  its  excretion  into  the  urine  occurs,  no  storage  of 
glycogen  occurs.  The  body  seems  quite  unable  to  metabolize  sugar  or  to 
convert  it  into  glycogen  as  in  the  normal.  From  such  observations 
physiologists  have  long  coupled  glycogenesis  with  the  function  of  the 
pancreas.  This  association  depends  upon  an  internal  secretion  of  the 
islands  of  Langerhans.  Extracts  of  pancreas  have  from  time  to  time  been 
experimentally  prepared  that  seemed  to  lower  the  amount  of  sugar  lost 
during  glycosuria.  Such  preparations  have  peculiar  general  toxicity 
and  little  progress  has  resulted  in  explanation  of  the  mechanism  of  sugar 
control. 

However,  Banting  and  Best  have  very  recently,  1922,  announced 
brilliant  results  in  the  solution  of  this  problem.  They  demonstrated  that 
intravenous  injections  of  a  watery  extract  of  pancreas  in  which  the 
parenchyma  had  been  degenerated  by  previous  operative  procedure,  see 
discussion  of  internal  secreting  glands,  when  injected  into  the  circulation 
of  a  depancreatized  dog  quickly  lead  to  the  disappearance  of  the  excess 


470  METABOLISM,    NUTRITION,    AND    DIET 

of  sugar  from  the  blood  and  to  a  decrease  in  its  loss  in  the  urine.  In  later 
epoch  making  announcements  from  the  University  of  Toronto  laboratories 
it  was  shown  that  glycogen  was  actually  stored  in  the  diabetic  liver  under 
the  influence  of  this  pancreatic  extract.  These  observations  confirm  the 
hypothesis  that  glycogenesis  in  the  liver  and  body  tissues  is  dependent  on 
the  presence  of  a  hormone  from  the  pancreatic  gland. 

The  Destination  of  Glycogen. — The  chief  theories  concerning  the 
use  of  glycogen  in  the  organism  are  advanced  by  Bernard  and  by  Pavy. 
The  former  considers  glycogen  as  a  reserve  supply  of  carbohydrate.  When- 
ever the  glycogen  of  the  blood  is  reduced  below  the  normal  level,  i.e.,  about 
o .  i  to  o .  1 5  per  cent. ,  there  is  a  conversion  of  glycogen  into  sugar.  The  sugar 
enters  the  blood  and  passes  to  the  tissues  where  its  oxidation  is  a  source  of 
energy.  Pavy  considers  glycogen  to  be  a  stage  in  the  synthesis  of  carbo- 
hydrate into  fat  and  protein.  Bernard's  theory  is  more  generally  accepted. 
It  explains  more  satisfactorily  why  the  sugar  content  of  the  blood  is  so  con- 
stant. The  conversion  of  glycogen  to  sugar  takes  place  by  the  action  of  an 
intracellular  ferment  in  the  glycogenic  cells.  Such  an  enzyme  has  been  iso- 
lated for  the  liver.  It  is  this  enzyme  that  converts  the  liver  glycogen  to  dex- 
trose after  death,  and  which  is  destroyed  by  boiling  in  the  usual  process  of 
isolating  glycogen  from  the  liver  or  other  tissues. 

Glycemia  and  Glycosuria. — Sugar  may  be  present  to  excess  not  only 
in  the  hepatic  veins,  but  in  the  systemic  blood.  When  such  is  the  case, 
the  sugar  is  excreted  by  the  kidneys,  and  appears  in  variable  quantities  in 
the  urine.  This  condition  is  known  as  glycosuria. 

Glycemia  and  glycosuria  may  occur  in  the  normal  animal  or  man 
during  the  absorption  of  the  products  of  digestion  following  a  meal  rich  in 
carbohydrates.  The  concentration  of  sugars  in  the  blood  occurs  because 
of  the  greater  rate  of  intake  than  rate  of  glycogen  storage.  This  leads 
to  a  digestion  glycemia  and  glycosuria.  However,  a  more  permanent 
glycosuria  may  be  produced  experimentally.  The  operative  removal  of 
the  pancreas  as  related  above,  leads  to  an  immediate  glycemia  and 
glycosuria.  This  condition  is  known  as  pancreatic  diabetes.  In  medicine 
it  forms  one  of  the  most  difficult  and  at  the  same  time  interesting  classes  of 
clinical  patients.  Puncture  of  the  medulla  in  the  region  of  the  vaso- 
motor  center  will  also  produce  diabetes.  In  fact,  any  sharp  disturbance 
of  the  central  nervous  system  may  be  followed  by  a  corresponding  dis- 
turbance in  sugar  metabolism.  The  administration  of  drugs,  such  as 
phloridzin,  strychnine,  glycosides,  morphine,  adrenalin,  amyl  nitrite,  or 
reduction  of  oxygen  or  excess  of  carbon  dioxide  may  be  followed  by  dis- 
charge of  glycogen  from  the  liver  store  houses  and  by  glycosuria.  Some  of 
these  agencies  react  on  the  central  nervous  system  influencing  either 
hormone  production  or  liver  activity  itself.  Other  agencies  like  adrenalin 
stimulate  the  peripheral  hepatic  nervous  mechanism.  In  any  event,  the 
normal  cycle  of  carbohydrate  metabolism  is  upset. 


THE  INFLUENCE  OF  MINERALS 


471 


THE  INFLUENCE  OF  MINERALS,  OF  FASTING,  ETC. 

Mineral  Matters,  Water,  Etc. — The  chief  mineral  constituents  of 
the  foods  are  sodium,  potassium,  calcium,  magnesium,  and  iron,  together 
with  chlorine,  sulphur,  and  phosphorus.  The  inorganic  substances  are  not 
a  source  of  heat.  They  may  supply  a  certain  amount  of  energy,  as  osmotic 
energy,  but  this  is  of  no  significance  as  compared  with  their  influence  on  the 
metabolism  of  organic  substances.  An  animal  fed  on  a  normal  food  de- 
prived of  the  mineral  constituents  survives  only  a  few  weeks  at  most. 

The  amount  of  mineral  matter  in  the  tissues  of  the  human  body,  exclusive 
of  the  skeletal  parts,  is  about  one  per  cent.  It  is  safe  to  say  that  this  is 
chiefly  in  complex  organic  combination  in  the  body.  The  daily  quantity 
excreted  is  about  twenty  to  thirty  grams.  This  quantity  enters  the  body  in 
the  food,  chiefly  in  combination  with  complex  compounds.  It  is  a  question 
as  to  what  per  cent,  of  organic  salts,  like  the  calcium,  the  phosphates,  and 
the  iron,  is  available  when  taken  into  the  body  in  inorganic  form. 

We  have  discussed  in  previous  chapters  the  role  of  certain  salts  in  their 
influence  on  metabolism;  for  example,  of  sodium,  potassium,  calcium,  iron, 
etc.  Foods  like  milk  and  eggs  are  especially  rich  in  calcium  and  phosphorus 
and  are  particularly  desirable  for  young  children,  the  former  for  its  influence 
on  the  growth  of  the  skeleton,  the  latter  for  the  same  reason  and  as  a  stimu- 
lator of  growth  of  protoplasm  in  general.  Lack  of  mineral  constituents, 
especially  calcium  compounds,  in  food  shows  its  influence  on  metabolism 
in  the  disease  known  as  rickets. 

NUTRITION  EXPERIMENT  IN  FIVE-MONTHS-OLD  PIGS.      (E.  B.  FORBES.) 


Per  cent,  gain   in    certain 

tissues   corresponding   to 

Per  cent,  gain 

i    per  cent,   gain   in  live 

in  live 

weight. 

Rations. 

weight  in 

60    days' 

feeding. 

Psoas 

Ash  of 

Thick- 

muscle. 

humer- 

ness    of 

us. 

back  fat. 

Hominy;  blood  flour;  bran  extract. 

69.  i 

.81 

•59 

.64 

(Phosphorus  mostly  as  phytin.) 

Hominy;  blood  flour;  bone  flour. 

61  .0 

.61 

.72 

.82 

(Phosphorus  mostly  as  tricalcic 

phosphate)  . 

Hominy  ;  blood  flour.      (Low  phos- 

41 .6 

.72 

.08 

i.o4 

phorus  ration). 

472 


METABOLISM,    NUTRITION,    AND   DIET 


Numerous  imvestigations  are  in  progress  which  may  demonstrate  more 
fully  the  specific  influence  of  phosphorus  on  animal  nutrition  and  on 
growth.  Tunnicliff  has  demonstrated  that  an  increase  of  the  phosphorus 
content  of  the  food  of  children,  if  given  in  complex  organic  form,  increases 
the  efficiency  of  the  metabolism  of  nitrogen  by  as  much  as  10  per  cent. 
If  given  to  children  as  calcium  phosphate  it  has  no  beneficial  influence 
in  this  regard.  Forbes,  in  his  experiments  on  the  nutrition  of  pigs,  shows 
that  the  individuals  fed  with  food  to  which  phosphorus  was  added,  as  ground 
fresh  bone,  grew  larger  and  stronger  skeletons,  but  that  the  presence  of 
organic  phosphorous  (phytin)  led  to  the  greatest  general  growth. 

PERCENTAGE  OP  PHOSPHORIC  ACID  (P2O6)  IN  SOME  FRESH  FOODS.     (QUOTED 

FROM   GlRARD,   BY  HuTCHINSON,   IN   "FOOD  AND  DIETETICS.") 


Vegetable.  Per  cent. 

Carrot 0.036 

Turnip o .  o  58 

Cabbage o .  089 

Potato o .  140 

Chestnuts o .  200 

Barley  meal o .  230 


Animal.  Per  cent. 

Pork o.i 60 

Milk 0.220 

Beef 0.285 

Eggs 0.337 

White  cheese °-374 

Mutton 0.425 


Salts  in  the  body  not  only  take  part  in  the  reactions  themselves,  but  they 
stimulate  in  other  substances  reactions  that  are  of  incalculable  benefit  to 
the  body. 

The  necessity  for  the  taking  of  water  in  order  to  balance  the  daily  ex- 
cretion, is  sufficiently  obvious.  Man  will  live  only  a  few  days  if  deprived  of 
water. 

Effects  of  Deprivation  of  Food. — The  animal  body  deprived  of  all 
food  dies  from  starvation  in  the  course  of  a  variable  time.  The  length  of 
time  that  any  given  animal  will  live  in  such  a  condition  depends  upon  many 
circumstances,  the  chief  of  which  are  the  nature  and  activity  of  the  metab- 
olism of  its  tissues. 

The  effect  of  starvation  on  the  lower  animals  is,  first  of  all,  as  might  be 
expected,  a  loss  of  weight.  The  loss  is  greatest  at  the  beginning  of  the  dep- 
rivation period,  but  afterward  decreases  to  a  level  from  which  it  does  not 
vary  much  day  by  day  until  death  ensues.  Chossat  found  that  the  ultimate 
proportional  loss  in  different  animals  experimented  on  was  almost  exactly 
the  same,  death  occurring  when  the  body  had  lost  40  per  cent,  of  its  original 
weight.  Different  parts  of  the  body  lose  weight  in  very  different  proportions. 
The  following  most  noteworthy  losses  are  taken,  in  round  numbers,  from 
the  table  given  by  Chossat: 


Fat  

Per  cent. 

Q? 

Liver  

Per  cent. 
52 

Blood       

7  ^ 

Muscles  

43 

Spleen 

7  I 

Nervous  tissues  

2 

Pancreas.  . 

6d 

EFFECTS    ON    DEPRIVATION    OF    FOOD 


473 


These  figures  are  in  practical  agreement  with  those  of  later  experimenters. 
They  show  that  the  chief  losses  are  sustained  by  the  adipose  tissue,  the 
muscles  and  glands.  The  nervous  structures  and  the  heart  are  maintained 
at  the  expense  of  the  other  tissues  and  show  but  little  change. 

The  effect  of  starvation  on  the  temperature  of  the  various  animals  ex- 
perimented on  by  Chossat  was  very  distinct.  For  some  time  the  variation 
in  the  daily  temperature  was  more  marked  than  its  absolute  and  continuous 
diminution,  the  daily  fluctuation  amounting  to  3°  C.  instead  of  0.5°  to  i°  C., 
60 


in 


2 

S 


DAYS  OF  FASTING 

FIG.  310. — The  Elimination  of  Urea  by  Dogs  during  Fasting. 
— •—  Following  2,500  grams  of  meat  in  the  food. 
-  Following  1,500  grams  of  meat  in  the  food. 
—  Following  minimal  amount  of  protein  in  the  food. 


(Voit.) 


as  in  health.  The  temperature  fell  very  rapidly  a  short  time  before  death, 
and  death  ensued  when  the  loss  had  amounted  to  about  16 . 2°  C.  It  has  been 
often  said,  and  with  truth,  that  death  by  starvation  is  really  death  from  want 
of  heat.  The  effect  of  the  application  of  external  warmth  to  animals  cold 
and  dying  from  starvation  is  more  effectual  in  reviving  them  than  the  ad- 
ministration of  food. 

The  symptoms  produced  by  starvation  in  the  human  subject  are  hunger, 
accompanied,  or  it  may  be  replaced,  by  pain,  referred  to  the  region  of  the 
stomach;  insatiable  thirst;  sleeplessness;  general  weakness,  and  emaciation. 
The  exhalations  both  from  the  lungs  and  from  the  skin  are  fetid,  indicating 


474  METABOLISM,    NUTRITION,    AND    DIET 

the  tendency  to  decomposition  which  belongs  to  badly  nourished  tissues; 
and  death  occurs  often  with  symptoms  of  nervous  disorder,  delirium,  or 
convulsions.  Death  commonly  occurs  within  from  six  to  ten  days  after 
total  deprivation  of  food.  This  period  may  be  considerably  prolonged  by 
taking  a  very  small  quantity  of  food,  or  even  by  water  alone.  The  cases 
so  frequently  related  of  survival  after  many  days  or  even  some  weeks  of 
abstinence  have  been  due  either  to  the  last- mentioned  circumstances,  or  to 
other  no  less  effectual  conditions  which  prevented  the  loss  of  heat  and 
moisture. 

During  the  starvation  period  the  excretions  diminish.  The  urea,  as  rep- 
resenting the  nitrogen,  falls  quickly  in  amount,  reaches  a  minimum  where  it 
remains  fairly  constant  for  several  days  before  death.  The  sulphates  and 
phosphates  undergo  much  the  same  type  of  reduction.  The  carbon  dioxide 
given  out  and  the  oxygen  taken  in  diminish.  The  feces  diminish  as  well  as 
the  bile.  It  is  highly  probable  that  the  greater  part  of  the  nitrogen  represents 
the  loss  of  weight  of  the  muscles. 

In  starvation,  then,  we  see  that  the  only  income  consists  of  water  and 
the  inspired  oxygen.  The  whole  of  the  energy  of  the  body  given  out  in  the 
form  of  heat  and  mechanical  labor  is  obtained  at  the  expense  of  its  own 
tissues,  there  being  as  a  result  a  constant  drain  of  the  nitrogen  and  carbon, 
not  to  mention  the  other  elements  of  which  the  tissues  are  composed.  It  is 
obvious  that  such  a  condition  cannot  be  endured  for  any  length  of  time. 

REQUISITES  OF  A  NORMAL  DIET. 

For  many  years  the  dictum  has  been  accepted  that  it  is  only  necessary 
that  a  normal  diet  should  be  made  of  the  various  classes  of  food  in  suffi- 
cient quantity  to  supply  to  the  normal  adult  animal  body  the  amount  of 
energy  that  is  lost,  due  to  the  daily  round  of  activity.  In  addition,  in 
young  and  growing  animals  an  excess  of  the  classes  of  foods  used  in  the 
construction  of  new  tissue  must  also  be  given.  No  doubt  these  desid- 
erata may  be  satisfied  by  many  combinations  of  foods,  and  it  would  be  un- 
reasonable to  expect  the  diet  of  every  adult  to  be  the  same.  The  age,  sex, 
strength,  and  circumstances  surrounding  each  individual  must  ultimately 
determine  what  he  takes  as  food.  A  dinner  of  bread  and  cheese  with  an 
onion  contains  all  the  requisites  for  a  meal,  but  such  diet  would  be  suitable 
only  for  those  possessing  strong  digestive  powers.  It  is  a  well-known  fact 
that  the  diet  of  the  continental  nations  differs  from  that  of  our  own  coun- 
try, and  the  diet  of  dwellers  in  cold  from  that  of  those  who  live  in  hot 
climates,  but  the  same  principle  underlies  all,  viz.,  the  replacement  of  the 
energy  losses  of  the  body  in  the  most  convenient  and  economical  way 
possible.  Any  one  in  active  work  requires  more  food  than  one  at  rest,  and 
growing  children  require  more  food  in  proportion  to  body  weight  than  adult 
men  and  women,  but  of  a  different  variety. 


REQUISITES    OF    A    NORMAL   DIET 


475 


The  chief  diet-scales  which  have  in  the  past  been  drawn  up  with  the 
object  of  supplying  the  proximate  principles  in  the  required  proportions 
are  given  in  the  table  below: 

STANDARD  DIETARIES. 


Author. 

Protein. 

Fat. 

Carbohy- 
drate. 

Calories 

Voit 

1  1  8     grams 

56     grams 

500     grams 

•3  o  <;  ? 

Rubner  
Moleschott       

127     grams 
I3°     grams 

52     grams 
40     grams 

509     grams 
5  50     grams 

3»°92 
3.  1  60 

Munk 

105     grams 

56     grams 

500     grams 

3.  022 

Wolff 

125     grams 

-I  ?     grams 

CAO     grams 

7  O7O 

Playfair 

1  1  o     grams 

ci     grams 

c-21     grams 

31  4.O 

Atwater 

12?     grams 

12?     grams 

A  co     grams 

3C2O 

Average  

121     grams 

59     grams 

510    grams 

3,135 

The  basis  of  computation  for  these  diets  is  to  supply  the  necessary  pro- 
tein nitrogen  consumed  by  the  tissues  of  the  body  first  of  all,  and  then 
to  supply  enough  potential  energy  to  balance  the  energy  expended  per 
day. 

The  amount  of  the  excreted  carbon  and  nitrogen  is  not  always  the  same. 
It  has  been  proven  possible,  for  example,  to  subsist  on  a  diet  containing 
9  or  10  grams  of  protein  nitrogen  and  200  grams  of  carbon  in  the  form  of 
carbohydrates  and  fats  per  diem,  the  ordinary  diet  for  needle-women  in 
London,  and  the  average  of  the  cotton  operatives  in  Lancashire  during  the 
famine  of  1862.  The  amount  of  these  elements  excreted  falls  to  figures  cor- 
responding to  such  an  income.  Of  course,  upon  such  a  diet  the  metabolism 
is  low,  and  persistent  physical  weakness  must  be  the  result,  probably  from 
insufficient  carbon.  The  9  or  10  grams  of  nitrogen  in  such  a  semi-starva- 
tion diet  would  be  equivalent  to  58.5  to  65  grams  of  protein,  whereas  the 
amount  of  protein  in  some  diets  may  be  as  high  as  1 50  and  more  grams  per 
day.  Chittenden's  nutritional  experiments,  so  often  referred  to  in  these 
pages,  have  proven  that  adult  men  can  subsist  in  nitrogenous  equilibrium 
and  do  vigorous  work  and  maintain  good  health,  on  a  protein  diet  below 
that  given  in  the  above  example,  i.e.,  on  6  to  10  grams  of  nitrogen.  In  such 
diets  a  plentiful  supply  of  carbohydrates  is  permitted,  but  the  caloric 
value  of  the  diet  is  less  than  those  in  the  table  above. 

Not  only  the  proteins  but  also  the  fats  may  vary.  The  amount  may  be 
as  low  as  35  grams  and  as  high  as  125  grams  The  carbohydrates  may 
vary  from  200  grams  to  500  grams  and  upward.  Sometimes,  with  a  small 
proportion  of  fats,  the  carbohydrates  may  be  correspondingly  increased 
to  make  up  the  necessary  quantity  of  available  potential  energy. 


METABOLISM,    NUTRITION,    AND   DIET 

From  the  data  on  page  342,  it  is  possible  to  form  various  diet-scales 
which  shall  supply  the  needs  of  different  conditions  of  growth  and  decay 
of  the  body.  In  an  economical  daily  ration  consisting  of  meat,  bread, 
butter,  cheese,  potatoes  and  vegetables  one  might  supply  the  requisite 
amount  of  protein  nitrogen  as  follows: 

N.  Proteins. 

340  grams  lean  uncooked  meat 10.0  grams  62.5  grams 

600  grams  bread 6.0  grams  37.5  grams 

90  grams  butter 0.5  grams  3.1  grams 

28  grams  cheese 1.5  grams  9.4  grams 

225  grams  potatoes  \     i . o  grams  6.2  grams 

225  grams  carrots.    / 

19.0  grams  118.7  grams 

The  30  grams  of  salts  necessary  to  replenish  the  daily  loss  by  excretion 
in  the  urine  are  contained  in  the  meat  16  grams,  the  bread  12  grams,  and 
vegetables  about  4  grams. 

The  fluid  should  consist  of  about  2,500  to  2,800  grams,  and  might  be 
given  as  water,  with  or  without  tea,  coffee,  or  cacao,  which  are  chiefly 
stimulants. 

The  Energy  Requirements  of  the  Body.— The  food  must  not  only 
make  up  for  the  substances  eliminated  from  the  body  but  must  also  supply 
the  potential  energy  of  heat  and  motion  set  free  in  the  living  body.  The 
amount  of  heat  is  measured  in  terms  of  calories,  or  more  often  in  large 
Calories.  The  work  energy  may  be  expressed  in  gram-centimeters,  or  in 
kilogrammeters.  Since  one  calorie  of  heat  is  the  equivalent  of  42,670  gram- 
centimeters  of  work,  the  two  units  may  be  computed  interchangeably. 

The  source  of  the  heat  and  work  energy  which  is  produced  in  the  body  is 
from  the  metabolic  changes  of  the  tissues,  the  chief  part  of  which  is  in  the 
nature  of  oxidation,  since  it  may  be  supposed  that  the  oxygen  of  the  atmos- 
phere taken  into  the  system  is  ultimately  combined  with  carbon  and  hydro- 
gen. Any  change,  indeed,  which  occurs  in  the  protoplasm  of  the  tissues, 
resulting  in  an  exhibition  of  protoplasmic  function,  is  attended  by  the  evolu- 
tion of  heat  and  the  formation  of  carbon  dioxide  and  water.  The  more  act- 
ive the  changes  the  greater  is  the  amount  of  heat  produced.  In  order  that 
the  protoplasm  may  perform  its  functions,  the  waste  of  its  own  destructive 
metabolism  must  be  repaired  by  the  due  supply  of  food  material  to  be  built 
up  in  some  way  into  the  protoplasmic  molecule.  In  the  tissues,  as  we  have 
several  times  remarked,  two  processes  are  continually  going  on:  the  building 
up  of  the  protoplasm  from  the  food,  anabolism,  which  is  not  accompanied 
by  the  evolution  of  heat;  and  the  oxidation  of  the  protoplasmic  materials, 
katabolism,  resulting  in  the  production  of  energy,  by  which  heat  is  set  free. 
Food  is  therefore  necessary  for  the  production  of  heat.  It  is  not  neccessary 
to  assume  that  the  combustion  processes,  indeed,  are  as  simple  as  the  bare 
statement  of  the  fact  might  seem  to  indicate.  But  complicated  as  the  vari- 


THE  ENERGY  REQUIREMENTS  OF  THE  BODY         477 

ous  stages  may  be,  the  ultimate  result  is  as  simple  as  in  ordinary  combustion 
outside  the  body,  and  the  products  are  the  same. 

This  view,  that  the  maintenance  of  the  temperature  of  the  living  body 
depends  on  continual  chemical  change,  chiefly  by  oxidation  of  combustible 
materials  in  the  tissues  or  by  the  tissues,  has  long  been  established.  The 
quantity  of  carbon  and  hydrogen  supplied  as  food,  chiefly  in  the  form  of 
carbohydrates  and  fats  which,  in  a  given  time,  unites  in  the  body  with 
oxygen,  is  sufficient  to  account  for  the  amount  of  heat  generated  in  the 
animal  within  the  same  period,  page  454.  This  amount  is  capable  of 
maintaining  the  temperature  of  the  body  at  from  36.8°  to  38.7°C.,  not- 
withstanding a  large  loss  by  radiation  and  evaporation.  This  estimation 
depends  upon  the  chemical  axiom  that  when  a  body  undergoes  a  chemical 
change  the  amount  of  energy  set  free  is  the  same,  supposing  the  resulting 
products  are  the  same,  whether  the  change  takes  place  suddenly  or  gradu- 
ally. If  a  certain  number  of  grams  of  different  substances  are  introduced 
as  food,  and  if  they  undergo  complete  oxidation,  the  amount  of  kinetic 
energy,  as  shown  in  the  amount  of  heat  and  mechanical  work,  is  the  same 
as  would  be  developed  if  the  same  bodies  were  completely  oxidized  outside 
the  body.  If  one  gram  of  fat  be  taken  into  the  body  and  is  completely 
oxidized,  resulting  in  the  production  of  a  definite  amount  of  carbon  dioxide 
and  water,  it  may  be  supposed  to  have  produced  the  same  amount  of  heat 
as  it  would  have  produced  outside  the  body.  In  the  case  of  protein  food 
it  is  a  little  different,  since  it  is  never  completely  oxidized  within  the  body, 
but  may  be  supposed  to  give  rise  to  a  definite  amount  of  urea  and  other 
lower  nitrogenous  compounds  not  completely  oxidized  in  the  body.  In 
this  case  the  gram  of  protein  may  be  considered  to  liberate  the  same 
amount  of  heat  as  the  protein  would  outside  the  body  minus  the  amount 
which  would  be  obtained  from  the  complete  oxidation  of  the  resulting 
urea,  etc. 

The  actual  amount  of  heat  produced  per  diem  has  been  experimentally 
ascertained  in  the  case  of  man  and  animals  by  the  aid  of  an  apparatus,  the 
calorimeter.  An  animal  is  enclosed  in  a  metal  cage  completely  contained 
in  a  second  cage  containing  water.  Air  is  let  into  and  out  of  the  inner  box 
by  means  of  metal  tubes  so  arranged  that  the  inlet  tubes  maintain  a  con- 
stant temperature  and  the  outlet  tubes  pass  through  water  between  the 
two  chambers.  The  heat  given  out  by  the  animal  warms  the  water  in 
the  outside  box,  and  may  be  estimated  by  the  rise  of  its  temperature,  the 
amount  of  which  is  known.  At  the  same  time  the  carbon  dioxide  output 
is  measured. 

The  rate  of  human  metabolism  has  come  to  be  of  great  importance  in 
clinical  diagnosis  of  nutritional  states.  The  direct  methods  are  too  in- 
volved for  rapid  testing.  Indirect  determinations  are  quite  accurate 
enough  for  such  purposes.  The  indirect  method  rests  on  measurement 


478 


METABOLISM,    NUTRITION,    AND    DIET 


of  the  rate  of  oxygen  consumption,  or  carbon  dioxide  output,  or  both,  as 
an  index  of  heat  production.  The  rate  of  heat  production  and  of  heat 
loss  must,  of  course,  be  in  balance  in  an  animal  of  constant  temperature,  as 
in  man.  For  this  reason  the  rate  of  metabolism  varies  not  with  mass, 
as  we  would  expect,  but  with  the  surface  area.  In  man  the  unit  used  is 
expressed  as  calories  per  square  meter  of  surface  per  hour.  Measured 
at  the  minimal  level  this  varies  according  to  the  following  averages. 

TABLE  OF  BASAL  METABOLIC  RATES. 

Rate  of  Minimal  Metabolism  in  Calories  per  Square  Meter  per  Hour,  Aub  and  DuBois, 
Ages  12-13  from  DuBois,  Logs,  from  Boothby  and  Sandiford. 


Age  in  years. 

Males. 

Log.  cals. 

Females  t             Log.  cals. 

14-16 

46.0 

6,628 

43-o 

6,335 

16-18 

43-o 

6,335 

40.0 

6,021 

18-20 

41  .0 

6,128 

38.0 

5,798 

20-30 

39-5 

5,966 

37-0 

5,682 

30-40 

39-5 

5,966 

36.5 

5,623 

40-50 

38-5 

5,855 

36.0 

5,563 

50-60 

37-5 

5,740 

35-o 

5,44i 

60-70 

36.5 

5,623 

34-o 

5,315 

70-80 

35-5 

5,502 

33-o 

5,185 

The  surface  area  for  man  is  computed  from  the  formula  of  DuBois  in 


ad 

190 
180 

|l70 

I.  i 

^IfiD 

0          30          40         50         60          70         80         90         100         1 

0 

200 

190 
180 
170 
160 
150 
140 
130 
120 
110 

100 
0 

s 

\ 

N^ 

N 

\ 

\ 

V.o 

x 

N2-I 

x 

*ai 

X 

23 

"X 

a 

\ 

\ 

\ 

\ 

\ 

M8 

X 

X 

X 

\ 

s^ 

~ 

X, 

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\ 

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V 

\ 

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\ 

\ 

x 

\_ 

X 

Sj 

-^ 

\ 

\ 

\ 

x 

\ 

\ 

\ 

\ 

\ 

N 

X 

X 

^ 

Zitn 

10 

\ 

\ 

\ 

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s 

\ 

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x 

\ 

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v» 

If) 

2Q 

o 

^140 

§30 

120 

no 

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X 

x 

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X, 

>r 

% 

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\ 

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rib 

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\ 

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\^ 

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^ 

^ 

'~20         30          40         50         60         70         80         90         100         1 
WEIGHT-KILOGRAMS 

FIG.  3ioa. — Height-weight  Chart. 

which  the  area  is  in  square  meters,  the  weight  in  kilograms,  and  the  height 
in  centimeters. 


THE  ENERGY  REQUIREMENTS  OF  THE  BODY         479 

Area  =  Weight0-425  X  Height0-725  X  Factor    71.84. 

The  area  is  more  quickly  determined  from  the  height-weight-graph 
computed  also  by  DuBois  and  DuBois. 

The  amount  of  heat  evolved  by  the  oxidation  of  various  food  stuffs  has 
been  carefully  measured  by  numerous  observers;  the  figures  calculated  by 
Rubner  are  perhaps  most  satisfactory. 

HEAT  VALUE  TO  THE  BODY. 

i  gram  carbohydrate 4.1  Calories. 

i  gram  fat 9.3  Calories. 

i  gram  protein 4.1  Calories. 

One  gram  of  dry  protein  has  a  total  heat  value  of  5.754  Calories  (Rubner), 
hence  it  is  obvious  that  protein  is  not  completely  oxidized  by  the  body.  Each 
gram  of  protein  yields  at  least  one-third  of  a  gram  of  urea,  which  has  a  heat 
value  of  2.5  Calories  per  gram. 

Atwater  and  Benedict  have  checked  the  energy  value  of  the  foods 
actually  consumed  against  the  actual  liberation  of  heat  and  work  energy  of 
the  human  body.  They  find  a  wonderfully  close  agreement  both  for 
periods  of  rest  and  for  periods  of  work.  Atwater's  estimate  for  the  energy 
needs  of  man  are  as  follows : 


Man  without  muscular  work 2,700   Calories. 

Man  with  light  muscular  work 3,000   Calories. 

Man  with  moderate  muscular  work 3,5°°   Calories. 

Man  with  severe  muscular  work 4, 500   Calories. 

The  daily  output  of  energy  for  the  adult  man  is  as  follows: 

Kilogrammeters.  Calories 

Work  of  heart  per  day 88,000 

Work  of  respiratory  muscle 14,000 

Eight  hours'  active  work 213,344 


315,334  or  743 

Amount  of  heat  produced  in  24  hours 1,582,700  or  3»724 


1,898,034  or  4,467 

This  estimate  is  relatively  high  for  ordinary  activity  as  determined  by 
Atwater  and  others.  It  is  indeed  more  energy  than  the  standard  diets  in 
the  table  given  on  page  450  will  yield  to  the  body.  For  example,  Voit's 
diet  yields  3,055  Calories,  and  the  average  of  the  table  is  only  3,125 
Calories. 


480  METABOLISM,    NUTRITION,   AND    DIET 

FOOD  SUPPLEMENTS,  THE  VITAMINES. 

Numerous  workers  have  shown  that  it  is  not  sufficient  to  supply  the 
body  with  adequate  proteins,  fats,  carbohydrates,  inorganic  salts  and  water 
meeting  the  standard  requirements  as  regards  the  caloric  value  of  foods. 
Formerly  these  facts  were  considered  as  physiologically  settled,  both  by 
the  medical  profession  and  by  those  interested  in  animal  nutrition.  In 
the  next  chapter  we  will  find  that  minute  quantities  of  substances  pro- 
duced by  the  various  ductless  glands  of  the  body  may  profoundly  influence 
the  process  of  metabolism.  In  like  manner,  observation  indicates  that  the 
foods  contain  similar  traces  of  material  supposedly  of  chemical  nature 
which  have  an  influence  on  metabolism  out  of  all  proportion  to  their 
quantity  or  mass.  Such  substances  have  been  given  the  name  of  vita- 
mines  by  Funk.  Most  food  sources,  such  as  meat,  milk,  fish,  vegetables 
and  cereals  contain  these  so-called  vitamines.  However,  the  amount  of 
vitamines  furnished  by  each  varies  widely. 

The  Nutritional  Diseases. — A  group  of  well  known  diseases  of  obscure 
etiology  have  been  investigated  on  a  large  scale  since  the  discoveries  of 
Funk.  At  the  present  time  the  diseases  of  xerophthalmia,  of  the  beri- 
beri type  of  neuritis,  and  the  scorbutic  diseases  are  definitely  attributed  to 
lack  of  corresponding  specific  vitamines  in  the  diet,  namely,  fat  soluble 
A,  water  soluble  B,  and  water  soluble  C  vitamines.  Other  factors  in  the 
food  are  necessary  to  its  complete  adequacy.  Of  these  it  is  evident  that  a 
proper  chemical  balance  among  the  inorganic  salts  of  the  diet,  as  well  as  a 
certain  bulk  and  quality  to  which  the  digestive  apparatus  of  the  particular 
species  of  animal  may  be  adapted,  plays  a  vital  part  in  the  food  sufficiency. 
Many  of  these  questions  are  not  yet  adequately  determined. 

Beri-beri  and  the  Antineuritic  Vitamine  B. — In  the  far  East  there 
is  prevalent  a  characteristic  nervous  disease  known  as  beri-beri.  It 
is  characterized  by  loss  of  nervous  function  which  comes  on  relatively 
suddenly  and  proceeds  to  a  state  of  motor  paralysis.  Beri-beri  has  been 
found  to  be  associated  with  the  use  of  an  excessive  rice  diet  and  is  prevalent 
in  rice  eating  countries.  Chickens,  pigeons,  and  pigs  which  are  relatively 
susceptible  to  a  neuritis  very  similar  to,  if  not  identical  with,  beri-beri 
quickly  recover  when  fed  on  rice  polishings  at  an  early  stage  of  the  disease. 
Such  experiments  have  led  to  the  conclusion  that  in  the  polishing  of  rice 
a  food  constituent  is  removed  that  is  absolutely  necessary  to  the  normal 
growth  of  the  nervous  system.  This  material  has  been  extracted  and 
chemically  examined  by  Funk  who  describes  it  as  an  organic  crystalline 
substance  melting  at  233°  C.,  chemically  resembling  pyrimidin.  Sub- 
stances like  the  rice  extract  which  cause  recovery  from  nutritional  diseases 
of  the  neuritic  type  have  been  given  the  name,  antineuritic  substances. 
Williams  prepared  an  oxypyridine  that  cured  neuritic  pigeons  in  doses 
of  one  milligram. 


THE    NORMAL   VITAMINES  481 

Scurvy  and  the  Antiscorbutic  Vitamine  C. — Scurvy  has  long  been  known 
to  be  due  to  the  inadequacies  of  a  restricted  diet,  for  example,  in  the  siege 
of  Paris,  in  long  ship  voyages,  etc.  The  disease  so  prevalent  in  the 
southern  part  of  the  United  States,  and  especially  in  the  Mississippi 
Valley,  known  as  pellagra  seems  to  be  in  the  same  category.  Through  the 
work  of  Voegtlin,  Koch  and  Sullivan  it  is  found  that  solutions  of  flour, 
corn  meal  and  vegetables  contain  a  substance  influencing  metabolism. 
This  has  been  tested  on  mice,  and  on  man  in  the  pellagra  hospitals. 
The  active  principle  is  permanent  in  acid  but  destroyed  in  the  alkaline 
solution.  For  example,  breads  that  have  been  leavened  by  soda  have  lost 
this  principle.  Soda  breads  are  detrimental  to  health  not  because  of  the 
manner  of  their  cooking  with  excess  of  oil,  but  rather  because  of  the 
destruction  of  the  vitamines  by  the  alkalinity  of  the  soda.  This  active 
type  of  vitamine  is  called  an  antiscorbutic  substance. 

Koch  and  Voegtlin  have  studied  the  central  nervous  system  of  man 
dying  of  pellagra,  also  the  nervous  system  of  monkeys  fed  on  inadequate 
diets.  The  results  show  a  loss  of  lipoids,  a  tendency  to  a  decrease  in  the 
proportion  of  proteins  and  an  increase  in  the  water  content.  There  is  a 
decrease  in  the  cerebrosides,  phosphatides  and  sulphatides  with  a  relative 
decrease  of  cholesterol  in  the  cerebrum  and  an  increase  in  the  spinal 
cord.  There  is  an  increase  in  extractives,  especially  the  nitrogenous 
extractives.  In  general,  "the  spinal  cord  exhibits  most  striking  chemical 
changes,  a  fact  which  is  in  perfect  agreement  with  histological  obser- 
vations." The  antiscorbutic  substances  are  present  in  cabbage,  tomatoes 
and  the  citrous  fruits. 

The  Growth  Producing  and  Antirachitic  Vitamine  A. — Osborne  and 
Mendel  have  performed  numerous  feeding  experiments  on  rats  which 
tend  to  show  that  special  growth  stimulating  substances  are  present  in 
foods.  They  find,  for  example,  that  milk  when  its  natural  proteins  are 
substituted  by  excess  of  casein  will  not  support  growth  for  a  long  time. 
When  natural  milk  or  milk  albumins  are  resupplied,  growth  proceeds. 
In  the  growth  producing  food  mixtures,  if  the  lard  which  they  used  for 
fat  is  replaced  by  butter  fat,  growth  proceeds  more  rapidly.  It  would 
seem  that  a  fat  soluble  growth  stimulating  substance  is  present  in  the 
foods 

If  the  fat  soluble  A,  so  richly  present  in  butter  fats,  liver  fat,  etc.,  is 
deficient  in  the  diet  a  characteristic  malnutritional  disease  of  the  eye  lids, 
cornea,  etc.,  occurs  known  as  xerophthalmia.  This  disease  clears  up  when 
fats  containing  the  vitamine  A  are  resupplied. 

Source  of  the  Normal  Vitamines. — Voegtlin  has  emphasized  the  fact 
that  the  animal  body  can  not  manufacture  its  own  vitamines  from  vitamine 
free  food.  The  vitamines  in  eggs  and  milk,  perhaps  accounting  for  the 
31 


482 


METABOLISM,   NUTRITION,   AND    DIET 


relatively  low  content  of  milk  in  vitamines,  apparently  come  from  plant 
food  sources.  The  plant  can  build  up  the  vitamines  from  simpler 
compound. 

TABLE   SHOWING  THE  RELATIVE  RICHNESS  IN  ANTINEURITIC  AND  ANTISCORBUTIC 
VITAMINES  FROM  COMMON  FOOD  SOURCES.     (FROM  VOEGTLIN.) 


Antineuritic  Properties 

Antiscorbutic  Properties 

Relatively  rich 

Relatively  poor 

Relatively  rich 

Relatively  poor 

Brewers'  yeast 

Sterilized  milk 

Fresh  vegetables 

Dried  vegetables. 

Egg  yolk 

Sterilized  meat 

Fresh  fruit 

Dried  fruit 

Ox  heart 

Cabbage 

Raw  milk 

Sterilized  milk 

Milk  (fresh) 

Turnips 

Raw  meat 

Canned  meat 

Beef   and   other   fresh 

Carrots    and    other 

Cereals,  sprouting. 

Dried  cereals 

meats 

vegetables   of   this 

Pork  fat 

Fish 

type 

Starch 

Beans 

Highly  milled  cereals 

Molasses 

Peas 

Starch 

Corn  syrup 

Oats 

Molasses 

Barley 

Corn  syrup 

Wheat 

Corn 

Other  cereals 

Methods  of  milling  rice,  wheat  flour  and  corn  products  whereby  the 
outer  hull  of  the  grain  which  contains  the  major  portion  of  the  vitamines 
is  lost  should  undergo  vital  modification.  Both  the  kernel  of  the  grain 
and  the  bran  which  formerly  have  been  largely  discarded  as  human  food 
are  rich  in  vitamines.  They  should  be  retained  in  the  cereal  products. 
The  public  agitation  for  the  use  of  whole  wheat  and  whole  grain  products 
has  its  biological  justification  chiefly  in  the  conservation  of  the  natural 
vitamine  supplies.  It  is  also  evident  that  in  cooking  and  in  the  prepara- 
tion of  food  care  should  be  taken  not  to  destroy  the  normal  vitamines  by 
an  excessive  use  of  alkali.  Heat  in  moderate  degree  somewhat  reduces 
the  normal  vitamines.  When  such  a  reduction  is  unavoidable  then  the 
cooked  foods  should  be  supplemented  by  vegetables  and  citrous  fruits  in 
which  the  vitamines  are  preserved. 


THE  INFLUENCE  OF  THE  DUCTLESS  GLANDS  ON  METABOLISM. 

A  further  question  to  be  considered  is  the  relationship  between  the 
metabolism  of  the  tissues  and  the  products  of  the  metabolism  of  other 
tissues. 

It  is  well  known  that  all  tissues  elaborate  substances,  either  end  products 
or  pure  waste  products,  which  may  more  or  less  sharply  influence  the  reactions 
in  the  other  tissues  of  the  body.  Many  of  the  waste  products  are  far  from 


INFLUENCE    OF    THE    DUCTLESS   GLANDS    ON    METABOLISM 


483 


inert  in  this  sense,  though  little  physiological  consideration  is  given  to  this 
class  of  material  in  this  regard.  On  the  other  hand,  many  end  products  of 
the  metabolism  of  certain  tissue,  the  ductless  glands  in  particular,  are 
thrown  back  into  the  blood  stream  to  influence  the  reactions  of  other 
tissues,  just  as  do  blood  constituents  from  outside  sources.  The  tissues 
and  organs  have  become  adapted  to  reaction  in  the  presence  of  chemicals 
of  this  type  and  in  many  instances  the  relation  is  so  firmly  established  as  to 
become  an  interdependent  one.  Adaptations  have  taken  place  as  between 
the  tissue  and  its  special  environment  of  material  derived  from  its  neigh- 


FIG.  311. — Part  of  a  Section  of  the  Human  Thyroid,  a,  Fibrous  capsule;  b,  thyroid 
vesicles  filled  with,  e,  colloid  substances;  c,  supporting  fibrous  tissue;  d,  short  columnar  cells 
lining  vesicles;/,  arteries;  g,  veins  filled  with  blood;  h,  lymphatic  vessel  filled  with  colloid 
substance.  (S.  K.  Alcock.) 

boring  sources.  Such  substances  are  called  internal  secretions,  and  the 
active  principle,  either  isolated  and  identified  or  assumed,  goes  by  the 
name  hormone  (Starling).  A  hormone  may  be  defined,  therefore,  as  a 
chemical  substance  produced  by  one  tissue  or  organ  to  which  some  other 
portion  of  the  body  has  become  biologically  adapted  to  such  an  extent 
that  its  normal  function  can  proceed  only  under  the  influence  of  the 
substance.  Hormones  are  more  or  less  specific  in  character.  A  hormone 
for  one  tissue  acts  particularly  on  that  tissue  to  the  practical  exclusion 
of  all  others. 


484  METABOLISM,    NUTRITION,    AND   DIET 

The  more  important  organs  which,  according  to  our  present  views, 
produce  internal  secretions  are  as  follows:  thyroid,  parathyroid;  pituitary 
posterior  lobe,  pituitary  anterior  lobe;  thymus;  suprarenal  cortex,  supra- 
renal medulla  and  chromamne  tissues;  the  islands  of  Langerhans  of  the 
pancreas;  duodenal  mucosa;  liver;  kidney;  testicular  interstitial  tissue; 
ovary  interstitial  tissue  and  corpus  luteum;  placenta,  and  fetus. 

THE  FUNCTION  OF  THE  THYROIDS. 

The  Thyroid  and  Accessory  Thyroids. 

The  thyroid  glands  with  certain  structures  called  accessory  thyroids 
are  situated  in  the  neck  to  either  side  of  the  trachea.  The  gland  consists 
of  two  lobes,  one  on  each  side  of  the  trachea  extending  from  the  thyroid 
cartilage  to  the  level  of  the  clavicle  or  lower.  Often  the  lobes  are  con- 
nected across  the  mid  line  by  a  middle  bar  or  isthmus.  The  thyroid  is 
covered  by  the  muscles  of  the  neck  It  is  highly  vascular  and  varies 
greatly  in  size  especially  in  different  parts  of  the  country.  The  gland 
vessels  are  each  lines  with  a  single  layer  of  cubicle  cells  producing  different 
sized  follicles  filled  with  a  transparent  colloid  material  containing  nucleo 
albumen  and  the  specific  secretion  of  the  gland. 

The  accessory  thyroids  are  detached  portions  of  thyroid  tissue  in  the 
neighborhood  of  the  lateral  lobes.  They  are  not  different  in  function  from 
the  rest  of  the  gland. 

The  Function  of  the  Thyroids. 

The  thyroids  now  are  known  to  secrete  a  specific  chemical  hormone 
which  tremendously  influences  the  rate  of  metabolism  of  the  body  as  a 
whole.  This  secretion  is  produced  by  the  cells  of  the  follicle  and  diffuses 
out  into  the  lymph  and  blood  stream  on  the  one  hand  or  into  the  colloid 
material  on  the  other.  Ultimately,  however,  it  enters  the  blood  stream 
and  is  generally  distributed  throughout  the  body. 

It  has  been  a  long  and  difficult  task  to  determine  the  influence  of  the 
thyroid  upon  metabolic  processes.  This  is  due  in  part  to  the  fact  that  the 
early  work  did  not  distinguish  between  the  thyroids  and  parathyroids,  and 
in  the  later  work  it  has  been  found  extremely  difficult  to  eliminate  the 
parathyroids  without  injury  to  the  thyroids. 

The  present  view  is  that  the  thyroid  hormone  increases  the  rate 
of  metabolism  throughout  the  body.  This  it  does  largely  by  virtue  of  the 
fact  that  the  iodine  containing  compound  acts  as  a  deamidizer  thus 
hastening  nitrogen  elimination  and  the  process  dependent  upon  nitrogen- 
ous metabolism.  This  last  conception  rests  chiefly  on  the  work  of  Kendall, 


THE  THYROIDS  AND  ACCESSORY  THYROIDS          485 

who  has  further  purified  the  hormone  and  determined  its  chemical  struc- 
ture. He  applies  the  term  thyroxin  to  the  compound.  Thyroxin  is  a 
compound  containing  60  per  cent,  of  iodine  and  its  rate  of  production 
depends  upon  the  presence  of  iodine  in  the  intake  of  the  body  foods.  The 
determination  of  the  influence  of  the  thyroid  has  been  a  long  and  interest- 
ing process.  The  steps  have  been  progressive  and  cumulative.  The 
methods  used  have  been  four.  First,  the  study  of  metabolism  in  thyroi- 
dectomy.  Second,  the  grafting  of  thyroid.  Third,  the  feeding  of 
thyroid  tissue.  Fourth,  the  isolation  and  physiological  administration 
of  the  purified  thyroid  hormone.  The  specific  dynamic  action  of  the 
thyroid  hormone  is  of  such  definite  character  that  at  the  present  time  the 
chief  dependence  of  medical  men  for  the  determination  of  variations  in 
the  gland  function  is  on  scientific  measurement  of  the  metabolic  rate 
under  the  conditions  known  as  basal  metabolism. 

a.  Thyroidectomy. — It  was  noted  in  the  middle  of  the  last  century  that 
complete  removal  of  the  thyroid  glands  produced  a  marked  derangement  of 
the  functions  of  the  body  and  generally  resulted  in  death.     Later  studies 
have  shown  that  death  was  due  to  the  fact  that  the  parathyroids  were  also 
removed.     The  influence  which  is  now  attributed  strictly  to  the  removal  of 
thyroids  consists  of  a  condition  of  malnutrition  characterized  by  marked 
myxedema,  a  trouble  that  also  appears  in  certain  thyroid  diseases.     Failure 
of  the  hormone  or  hypothyroidism  leads  to  a  depression  of  the  neuro- 
muscular  tone,  and  malnutrition  of  the  vascular  and  lymphatic  systems. 
In  the  early  atrophy  of  this  gland  the  body  fails  to  develop  and  cretinism 
is  the  result. 

b.  Thyroid  Grafting. — If  a  portion  of  the  thyroid  gland  be  engrafted  in 
any  other  locality  in  an  animal  and  the  graft  be  allowed  to  grow  before  the 
thyroids  are  removed,  then  the  characteristic  symptoms  no  longer  develop. 
The  small  amount  of  grafted  tissue  is  able  to  produce  enough  of  the  thyroid 
hormone  to  supply  the  needs  of  the  body.     Successful  and  beneficial 
results  have  now  been  obtained  by  transplanting  thyroid  gland  in  man 
(McPherson) .     If  in  experimental  cases,  the  thyroid  graft  be  removed,  then 
the  usual  symptoms  immediately  appear. 

c.  Thyroid  Feeding. — If  the  thyroid  is  fed  either  as  the  purified  hormone  or 
merely  as  the  gland  itself,  it  seems  to  stimulate  certain  types  of  metabolism, 
as  indicated  by  an  increase  in  nitrogenous  wastes,  and  by  more  healthful 
reaction  of  the  nerve  tissue.     There  is  also  an  increased  oxidation  of  fats, 
a  property  that  has  brought  extracts  of  this  gland  into  extensive  but  more  or 
less  dangerous  use  as  an  antifat  remedy.     Thyroid  feeding  has  the  same 
effect  on  the  body  as  hvperthyroidism  characteristic  of  certain  types  of  enlarged 
or  hypertrophied  gland,  exophthalmic  goiter,  or  in  hypothyroidism  from  any 
cause,  extraneous  use  of  the  thyroid  hormone  in  any  form  antagonizes  the 
symptoms  of  malnutrition.     Even  following  thyroidectomy  if  the  gland  or  its 
extract  be  fed  by  way  of  the  mouth  the  symptoms  disappear  or  are  reduced. 


486 


METABOLISM,   NUTRITION,    AND   DIET 


The  Nature  of  the  Thyroid  Hormone. — The  active  principle  produced 
by  the  thyroid  gland  is  an  iodine  compound  as  was  first  shown  by  Baumann 
in  1895.  He  prepared  an  iodine-protein  compound  called  thyroiodinc, 
which  contained  as  much  as  9.3  per  cent,  of  iodine.  The  compound  was 
stable,  readily  soluble  in  dilute  alkalies,  and  did  not  decompose  with  high 
heat  or  when  the  gland  was  digested  in  strong  acid,  Kendel's  thyroxin 
we  now  know  to  be  the  pure  principle. 

Marine  who  has  extensively  studied  the  thyroid  finds  that  certain  types 
of  hyperplasia  are  associated  with  a  decreased  supply  of  the  active  iodine 
compound.  If  iodine  as  such  be  given  to  such  an  animal,  the  symptoms  are 
improved  and  the  iodine  compound  content  of  the  gland  is  increased. 

Interesting  biological  experiments  showing  the  influence  of  thyroid 
feeding  on  immature  animals  have  been  performed  by  Gudernatsch.  Dried 


FlG.  312. — The  influence  of  thyroid  feeding  on  metamorphosis  of  tadpole  larvae  of 
Bufo  vulgaris.  a,  original  size  June  5th.  b,  control,  d,  and  e,  thyroid  and  plants,  d, 
thyroid  given  three  days,  e,  thyroid  given  five  days,  taken  June  26th.  m,  control,  n,  fed 
thyroid  and  muscle,  o  fed  thyroid  and  plants,  p,  thyroid  and  hypophysis,  taken  July  6, 1912. 
(After  Gudernatsch.) 

thyroid  gland  fed  to  different  species  of  tadpoles  stopped  growth  and 
stimulated  differentiation.  Tadpole  larvae  no  matter  how  young  were  forced 
into  metamorphosis,  i.e.,  the  development  of  fore  and  hind  limbs  and  the 
assumption  of  the  adult  frog  form.  Gudernatsch  by  this  means  stimulated 
even  tiny  tadpoles  only  a  few  millimeters  long,  forcing  metamorphosis 
"within  eighteen  days  after  hatching,  while  normally  they  would  require 
ten  to  twelve  weeks  to  reach  such  a  stage.  The  results  of  this  premature 
metamorphosis  are  perfect  frogs  of  minute  size,  pygmy  frogs."  The  thyroid 
as  food  stops  tissue  growth  during  this  process.  "The  thyroid  possesses  a 
quality  that  stimulates  differentiation,  not  contained  in  any  other  food 
used."  The  influence  of  the  thyroid  is  somewhat  modified  by  the  food 
given  with  the  gland,  but  one  food  would  prevent  the  thyroid  influence, 
namely  thymus. 


PARATHYROIDS 


487 


Parathyroids. — There  are  smaller  glandular  structures  associated 
with  the  thyroids  on  each  side  known  as  the  parathyroids.  One  is  located 
at  the  level  of  the  lower  border  of  the  cricoid  cartilage  and  in  intimate  rela- 
tion to  the  posterior  border  of  the  lateral  lobe  of  the  thyroid.  The  other  is 
at  the  inferior  border  of  the  same  lobe.  In  the  dog  there  is  an  anterior  para- 
thyroid at  the  head  of  the  thyroid  and  an  external  imbedded  in  the  mass  of 
the  thyroid.  Their  structure  is  somewhat  different  from  that  of  the  thyroid, 
and  it  has  been  claimed  that  after  the  removal  of  the  thyroid,  the  parathyroid 
tissue  develops  a  structure  more  nearly  comparable  to  the  thyroid.  The 
parathyroids  bear  a  close  relation  anatomically  and  physiologically  to  the 
thyroids.  It  is  still  the  view  that  the  parathyroids  are  intimately  associated 
with  the  thyroids  in  the  construction  of  the  thyroid  compound,  and  Mendel 
has  found  iodine  in  the  parathyroids.  This  supports  the  view  that  the  para- 

Epith.  c. 


on.  tis. 


Coll. 


— A 


Epith.  c- 


Epith.  c.-l 


-•>.- 


Thy. 


Ves. 


'Coll. 
•Bid.  t. 


-Ves. 
Conn.  I 


Coll. 


FIG.  313. — Normal  parathyroid  tissue  of 
the  cat.     (Vincent  and  Jolly.) 


FIG.  314. — Parathyroid  of  the  cat 
after  the  thyroids  have  been  removed, 
showing  enlarged  vesicles  with  colloid. 
(Vincent  and  Jolly.) 


thyroids  prepare  the  iodine  compound  which  is  later  stored  in  the  thyroid. 
A  supplementary  view  by  Koch  is  presented  later. 

a.  Parathyroidectomy. — The  removal  of  the  parathyroids  is  more  dis- 
turbing, on  dogs  at  any  rate,  than  the  removal  of  the  thyroids.  Dogs  survive 
the  operation  only  a  short  time,  three  to  five  days.  They  show  deranged 
metabolism  and  quickly  develop  a  characteristic  disturbance  of  nerve 
muscular  control  to  which  the  name  of  tetany  has  been  applied.  Restless- 
ness, excitability,  muscular  tremors  which  in  the  later  period  pass  into  mild 
convulsions,  with  tetanic  rigor  and  exhaustion,  followed  by  death  is  the  cycle 
of  events  after  this  operation.  It  has  also  been  shown  that  the  removal  of  the 
parathyroids  in  the  dog  leads  to  an  increased  excretion  of  calcium  and  of 
ammonia  and  of  other  products  indicative  of  disturbed  metabolism.  The 


488  METABOLISM,    NUTRITION,    AND   DIET 

derangement  of  calcium  metabolism  suggests  the  thought  that  the  symptoms 
might  be  relieved  by  supplying  this  substance.  The  matter  is  presented 
in  the  following  words  of  Berkeley  and  Beebe:  "Probably  the  most  striking 
argument  in  favor  of  Macallum's  theory  in  respect  to  tetany  parathyreo- 
privus  is  that  it  may  be  relieved  almost  instantly  by  the  intravenous  injec- 
tion of  a  soluble  calcium  salt.  We  have  repeated  the  experiment  and  find 
that  intravenous  injections  of  a  soluble  calcium  salt  will  relieve  tetany  almost 
instantly;  intramuscular  injections  may  be  effective  after  a  period  of  thirty 
to  forty-five  minutes,  and  a  similar  period  is  required  for  the  beneficial 
effect  after  subcutaneous  injection.  The  effect  of  the  calcium  salt  when  given 
intravenously  to  an  animal  in  advanced  states  of  acute  tetany  is  one  of 
the  most  striking  in  the  range  of  physiological  experimentation." 

Koch  in  1913  studied  the  metabolism  of  dogs  after  parathyroidectomy, 
using  physiological  chemical  methods,  supplemented  by  histological  examina- 
tion. The  urine  of  these  dogs  contained  the  toxic  substances  methylguanidine 
and  other  guanidine  bases  in  large  quantities.  These  are  purin  compounds. 
The  examination  of  the  tissues  showed  extensive  chromatolysis,  degenerating 
epithelial  cells  in  the  intestinal  tract,  and  nuclear  disintegration  in  the  liver 
and  especially  the  kidney.  These  led  him  to  the  view  that  the  absence  of 
the  parathyroids  resulted  in  disturbance  of  nuclear  metabolism,  a  "  nuclear 
atrophy."  It  has  already  been  shown  that  nucleoproteins  are  richly  present 
in  the  parathyroids  and  their  extracts.  Injections  of  nucleoproteins  freshly 
prepared  also  relieve  the  symptoms  of  acute  tetany  in  dogs.  Berkeley 
and  Beebe  state:  "We  believe  that  the  essential  fact  in  the  production  of 
symptoms  following  complete  thyroparathyroidectomy  is  the  deranged 
metabolism  giving  rise  to  some  active  poison,  and  not  the  abnormal  excretion 
of  calcium  which  we  regard  as  an  accompanying  phenomenon,  perhaps  due 
in  part  to  the  starvation  cachexia  which  ensues  in  most  cases  if  the  animal 
is  tided  over  the  acute  condition."  This  view,  it  will  be  noted,  was  borne  out 
by  the  later  work  of  Koch. 

The  Suprarenal  Capsules  or  Adrenals. — These  are  two  flattened, 
more  or  less  triangular  or  cocked-hat  shaped  glandular  bodies,  resting  by 
their  lower  border  upon  the  upper  border  of  the  kidneys. 

The  gland  tissue  proper  consists  of  an  outside  firmer  cortical  portion, 
and  an  inside  soft  dark  medullary  portion,  figure  315. 

The  adrenals  are  very  abundantly  supplied  with  nerves,  chiefly  com- 
posed of  non-medullated  fibers.  These  fibers  are  derived  from  the  solar 
and  renal  plexuses  and  the  vagi,  but  the  method  of  their  termination  is 
unknown. 

A  vast  amount  of  information  has  been  given  concerning  the  function 
of  the  suprarenal  capsules  within  the  last  few  years  by  the  researches  of 
Oliver  and  Schafer,  Zyboulski,  Abel,  Aldrich,  Dakin,  Dale,  Elliot,  Cannon, 
and  others.  Brown-Sequard,  it  is  true,  showed  by  experiment  as  early 


THE    SUPRARENAL    CAPSULES    OR    ADRENALS 


489 


as  1856  that  removal  of  the  suprarenals  is  followed  by  the  death  of  the 
animal,  but  his  experiments  were  repeated  by  others  unfortunately  on 
less  susceptible  animals.  These  men 
did  not  obtain  the  same  results;  and 
it  was  concluded  that  the  suprarenal 
capsules  had  no  function,  or  at  least 
that  their  function  was  not  known.  l 
Death  was  preceded  in  the  case  of 
Brown-Sequard's  animals  by  symptoms 
somewhat  analogous  to  those  of  the  b 
disease  of  man  known  as  Addison's 
disease.  The  failures  to  reproduce  these 
symptoms  have  probably  resulted  from 
incomplete  removal  or  the  presence  of 
accessory  bodies.  Accessory  suprarenal- 
like  tissues,  chromofile  tissues,  are  com- 
monly present  in  animals.  Further, 
if  one  gland  is  removed,  the  other 
hypertrophies.  The  experiments  of  all 
recent  observers  confirm  the  original 
experiments  of  Brown-Sequard  and  show  c 
that  the  removal  of  the  suprarenal  cap- 
sules is  fatal. 

Oliver  and  Schafer  found  that  injec- 
tions of  suprarenal  extract  produced 
marked  effects  upon  the  muscular  layer 
of  the  arteries,  the  muscular  tissue  of 
the  heart,  and  the  skeletal  muscles. 
The  muscular  layer  of  the  arteries  is 
markedly  contracted,  causing  a  rise  of 
blood-pressure. 

The  contractions  of  the  heart  with 

its  nerves  intact  are  small  although  they 

j    T_    .LI     •      t  j  i*  FIG.     315. — Vertical     Section     of 

are  increased  both  in  force  and  ampli-   AdrenaL  */,  Capsule;  B,  cortex;  C, 

tude.      If    the    vagi    are    sectioned,    the    medulla;  a,  glomerular  zone;  b,  fasci- 
i       •         j  T  j     ,  v       cular  zone;  c,  reticular  zone;  v,  vein 

slowing  does  not  occur.     Instead,  the  in  medulla;    '(Merkei-Henle.) 

heart   rate   is   accelerated  through  the 

accelerator  nerves,  the  movements  of  the  alimentary  canal  are  de- 
creased, the  splanchnic  vascular  region  is  sharply  constricted,  the 
urogenital  musculature  is  stimulated  to  contraction,  and  the  pupils  are 
dilated.  In  fact,  Elliot  and  others  after  him  have  shown  that  all  those 
structures  which  are  stimulated  or  inhibited  by  the  action  of  sympathetic 
mechanisms  are  stimulated  by  the  active  principle  of  this  gland  and  in 


49°  METABOLISM,    NUTRITION,    AND   DIET 

the  same  manner.  The  conclusion  follows  that  the  active  principle, 
epinephrin,  specifically  stimulates  the  sympathetic  and  autonomic 
mechanisms.  The  point  of  action  is  the  nerve  endings  in  the  peripheral 
tissue  and  the  reaction  is  sympathomimetic. 

a.  The  Nature  of  the  Active  Principle  of  the  Suprarenal  Gland. — Early 
workers,  notably  Oliver  and  Schafer,  studied  this  gland  -by  extracts  of  its 
dried  tissue.  Thus  they  stated  that  less  than  yT  J-^ gram  (%%-$ grain)  of  the 


FIG.  316. — Injection  of  Suprarenal  Extract.     Effect  upon  the  heart,  limb,  spleen  and 
blood-pressure,  after  section  of  cord  and  vagi.     (Oliver  and  Schafer.) 

desiccated  gland  is  sufficient  to  produce  an  effect  upon  the  heart  arid  arteries 
of  an  adult  man. 

It  is  a  curious  fact  that  only  extracts  of  the  medullary  portion  of  the  gland 
are  active.  It  has  been  further  shown,  by  Christiani  and  others,  that  if  only 
small  portions  of  the  medulla  remain,  the  animal  operated  upon  survives; 
while  if  all  the  medullary  substance  be  removed,  even  though  large  portions 
of  the  cortex  remain,  the  animal  invariably  dies. 

Abel  has  succeeded  in  separating  the  blood-pressure-raising  constituent 
of  the  gland  extract,  and  calls  it  epinephrin,  C10H13NO3  JH2O.  Takamine 
isolated  an  active  principle  to  which  he  assigned  the  formula  C9H13NO3, 
and  the  name  adrenalin.  This  product  is  now  known  to  be  an  amino- 
alcohol  with  a  pyrocatechin  base  as  follows: 


THE  INTERNAL  SECRETION  OF  THE  PANCREAS        491 

HO/\CH.  OH.  CH2NH.  CH, 


This  compound  is  the  active  principle  or  hormone  of  the  epinephros.  Epi- 
nephrin  is  now  prepared  synthetically  and  commercially,  and  is  an  important 
and  useful  medicinal  product. 

b.  Secretory  Nerves  for  the  Adrenals. — Dreyer  was  first  to  give  evidence 
that  the  products  of  this  gland  are  discharged  into  the  blood  of  the  adrenal 
vein  in  increased  quantity  on  splanchnic  stimulation.     Cannon  has  exten- 
sively studied  the  relation  of  the  secretion  of  epinephrin  to  the  autonomic 
reactions  of  the  body.     His  evidence  indicates  that  during  fright,  anger, 
etc.,  which  in  animals  is  accompanied  by  vigorous  activity  of  the  mechanisms 
innervated  through  the  automatic  system,  the  suprarenal  glands  are  strongly 
stimulated.     The  result  is  a  production  and  discharge  of  an  increased  amount 
of  epinephrin  into  the  circulation.     As  the  nervous  mechanisms  tend  to 
fatigue  the  increase  in  the  secretion  of  epinephrin  brings  about  a  hormone 
stimulation  supplementing  and  strengthening  the  nerve  control  during  this 
critical  time. 

This  gland  furnishes,  on  the  whole,  most  conclusive  and  typical  evidence 
of  the  presence  of  an  internal  secretion  that  is  absolutely  necessary  to  the 
normal  metabolism  of  other  organs. 

c.  Epinephrin  in  Relation  to  Carbohydrate  Metabolism. — The  action  of 
the  hormone,  epinephrin,  which  seems  so  specific  in  relation  to  the  function 
of  special  types  of  nerve  endings  is  bound  up  with  the  metabolism  of  carbo- 
hydrates.    Following  the  removal  of  the  suprarenal  glands  the  store  of  gly- 
cogen  in  the  liver  is  quickly  discharged  and  glycogenesis  is  no  longer  possible. 
This  has  been  shown  by  removing  the  suprarenals  in  rats  which  stand  the 
operation  more  readily  than  most  animals,  if  the  glands  are  removed  in 
successive  operations. 

The  Internal  Secretion  of  the  Pancreas. — Minkowski  and  von 
Mering  have  shown  that  total  extirpation  of  the  pancreas  is  followed  in  all 
cases  by  the  appearance  of  sugar  in  the  urine  in  the  course  of  a  few  hours. 
The  amount  of  sugar  which  appears  is  considerable,  from  5  to  10  per  cent. 
This  experimental  disease  is  accompanied  by  an  increase  in  the  quantity  of 
urine  and  by  abnormal  thirst  and  appetite.  It  proves  fatal  in  fifteen  days  or 
less.  These  results  are  obtained  only  when  the  entire  gland  or  more  than 
nine-tenths  of  it  have  been  removed.  If  one-tenth  or  more  of  the  gland  be 
left  behind,  sugar  appears  in  the  urine  when  carbohydrates  are  eaten,  but  not 
otherwise.  Nor  is  it  necessary  that  the  remaining  portion  of  the  gland  be  in 
its  normal  situation.  Successful  grafts  of  pancreas  under  the  skin  of  the 
abdomen  or  elsewhere  will  prevent  the  appearance  of  sugar  in  the  urine  and 
the  other  symptoms.  If,  however,  the  graft  be  subsequently  removed,  the 
sugar  in^the  urine  and  the  other  symptoms  reappear,  and  the  experimental 


49  2  METABOLISM,    NUTRITION,    AND    DIET 

disease  proceeds  to  a  rapidly  fatal  issue.  Opie  and  others  have  shown  also 
that  in  most  cases  of  diabetes  mellitus,  there  are  pathological  changes  in  the 
islands  of  Langerhans. 

The  symptoms  produced  by  total  extirpation  of  the  pancreas  do  not  de- 
pend upon  the  loss  of  the  pancreatic  juice  proper  to  the  organism.  This 
secretion  may  be  diverted  from  the  intestine  through  a  pancreatic  fistula 
without  the  production  of  diabetes.  Moreover,  Hedon  and  Thiroloix  have 
rendered  the  acini  of  the  gland  functionally  inactive,  and  ultimately  de- 
stroyed them,  by  the  injection  of  paraffin  or  other  substances  into  the  duct 
of  Wirsung,  without  the  supervention  of  diabetes. 

These  experiments  have  shown  that  the  ordinary  secreting  cells  degener- 
ate and  the  islands  of  Langerhans  increase  in  size,  leading  to  the  conclusion 
that  the  islands  are  the  structures  that  produce  a  special  internal  secretion 
which  influences  or  controls  carbohydrate  metabolism  in  the  body.  When 
the  dextrose  cannot  be  readily  burned  in  the  body,  the  content  of  this  sub- 
stance in  the  blood  is  increased  and  this  excess  is  eliminated  by  the  kidneys. 
Products  of  incomplete  oxidation  of  the  sugar  are  also  thrown  into  the  circu- 
lation and  eliminated  into  the  urine.  In  diabetic  cases  oxybutyric  acid, 
aceto-acetic  acid,  acetone,  etc.,  are  found  in  the  blood  and  urine.  Neither 
pancreatic  nor  muscle  extracts  have  the  power  of  burning  sugar. 

Banting  and  Best,  1922,  have  announced  in  the  Journal  of  Laboratory 
and  Clinical  Medicine  a  most  brilliant  demonstration  of  the  isolation  and 
function  of  an  active  internal  secretion  from  the  pancreas.  They  per- 
formed a  two  stage  operation  on  dogs,  the  first  operation  consisting  of 
ligation  of  pancreatic  ducts  to  induce  degeneration  of  the  pancreatic 
acini.  Ten  months  later  they  removed  this  degenerated  pancreas  and 
injected  watery  extracts  of  it  into  the  circulation  of  the  animal  after 
the  development  of  the  usual  pancreatic  diabetic  glycaemia.  The  blood 
sugar  rapidly  augmented  after  removal  of  the  pancreas,  from  .09  to  over 
.30  per  cent.  There  was  a  corresponding  glycosuria,  as  indicated  in 
figure  3160.  On  injecting  5  cubic  centimeters  of  a  four  day  old  extract 
of  the  degenerated  pancreas,  the  blood  sugar  dropped  from  .27  to  .15 
per  cent,  but  slowly  returned  to  the  higher  level.  Repeated  injections 
of  degenerated  pancreatic  extract  continued  to  depress  the  amount  of 
blood  sugar.  These  injections  greatly  reduced  the  associated  glycosuria, 
see  figure  3160.  Dr.  Macleod,  in  whose  laboratory  these  experiments 
were  performed,  has  announced  before  a  scientific  medical  assembly  the 
extension  of  these  observations.  In  his  laboratory  it  has  been  shown  that 
the  active  principle  is  easily  destroyed  by  the  digestion  enzymes  and  by 
heat.  The  active  principles  must  be  injected  into  the  blood  stream  to 
produce  its  physiological  effect.  Furthermore,  if  the  active  principle 
is  injected  into  the  blood  stream  along  with  an  excess  of  glucose  the  liver 


THE    KIDNEY 


493 


will  store  up  glycogen.  This  epoch  making  point  has  never  before  been 
demonstrated  in  experimental  pancreatic  diabetes.  Preliminary  tests  on 
the  human  disease  show  marked  improvements  in  the  physical  and  mental 
symptoms  with  a  reduction  of  glycaemia  and  glycosuria.  If,  as  the 
authors  express  the  hope,  quantity  production  of  the  active  internal 
secretion  of  the  pancreas  can  be  perfected,  it  will  give  for  the  first  time  a 
direct  method  of  supplementing  the  physiological  deficit  in  the  necessary 
internal  secretion  for  controlling  the  metabolism  of  carbohydrates  in  the 
human  body. 


AU6.4 


FIG.  316^. — (i)  5  cc.  4  day  old  extract  of  degenerated  pancreas.  (2)  5  cc.  extract 
of  liver.  (3)  5  cc.  extract  of  spleen.  (4)  5  cc.  extract  of  degenerated  pancreas. 
(Banting  and  Best.) 


The  Liver  as  an  Internal  Secreting  Organ. — That  the  liver  is  an  organ 
of  manifold  function  is  well  known.  The  secretion  of  bile,  the  glycogenic, 
lipogenic,  urea,  and  uric  acid  forming  functions  have  already  been  dis- 
cussed. The  glycogenic  function  as  a  mechanism  for  maintaining  a  com- 
paratively constant  supply  of  carbohydrate  in  the  blood  is  a  typical  internal 
secretory  process  though  the  secretion  is  in  this  case  not  specific  and  not  a 
typical  hormone.  Lipogenesis  is  of  the  same  class.  The  urea,  and  in 
birds  uric  acid,  are  waste  products.  The  liver  carries  on  these  functions 
by  the  aid  of  enzymes,  i.e.,  oxidases,  etc.  There  is  also  increasing  evidence 
that  the  liver  is  concerned  with  the  metabolism  of  the  amino  acids  though 
the  problem  is  not  yet  adequately  understood. 

The  Kidney. — The  proof  of  an  internal  secretion  by  the  kidney  is  not  so 
clear,  chiefly  because  of  the  disturbance  in  the  elimination  of  waste  pro- 
ducts when  this  organ  is  removed.  However,  the  convulsions  which 
follow  nephrectomy  are  said  to  be  diminished  in  intensity  by  the  use  of 
extracts  of  the  kidney.  It  is  claimed  therefore  that  the  kidney  produces  an 
internal  secretion  in  addition  to  its  primary  function  as  an  excretory 
organ. 


494 


METABOLISM,    NUTRITION,   AND   DIET 


The  Intestinal  Mucosa. — The  duodenal  mucosa  under  the  stimulating 
influence  of  hydrochloric  acid  produces  a  secretion,  secretin,  which  acts  as 
a  chemical  hormone  to  the  pancreas.  The  enzyme,  enterokinase,  is  activat- 
ing for  the  trypsinogen  of  the  pancreatic  juice  although  the  reaction  takes 
place  within  the  intestine. 

The  Thymus  Gland. — The  thymus  is  a  ductless  gland  which  is  of  rel- 
atively large  size  during  embryological  development.  It,  like  the  thyroids 


/i.  thffr.JST 


FIG.  317. — Diagram  representing  the  embryological  origin  from  the  branchial  organ  of  the 
thyroids,  parathyroids,  thymus,  and  posterior  branchial  body.     (Vincent.) 

and  parathyroids,  arises  from  the  invagination  of  the  epithelium  of  the  third 
and  fourth  branchial  arches.  Little  is  known  directly  of  the  functions  of  the 
thymus  in  the  human.  Certain  cases  of  excessive  lymphoid  development 
in  adults  are  associated  with  persistence  or  enlargement  of  the  thymus. 
Thyroidectomy  interferes  with  the  full  development  of  the  thymus  and  is 
associated  with  other  evidences  of  disturbed  nutrition,  i.e.,  muscular, 
nervous,  skeletal,  etc. 

Important  information  has  been  recently  yielded  by  the  biological 
studies  of  Gudernatsch  referred  to  in  the  discussion  of  the  thyroid  gland. 
This  observer  fed  dried  thymus  to  frog  tadpoles  with  the  result  that  growth 
was  strongly  stimulated,  whereas  metamorphosis  was  indefinitely  delayed. 
Tadpoles  with  ordinary  food  begin  the  differentiation  that  leads  to  metamor- 
phosis upon  reaching  a  certain  maximum  size  and  age.  The  thymus-fed 
tadpoles  continue  to  grow  beyond  the  normal  size  without  showing  signs  of 
differentiation.  Whereas  the  thyroid  stops  growth  and  stimulates  differ- 
entiation, the  thymus  favors  growth  and  seems  to  prevent  differentiation. 

The  Pituitary  Body. — This  structure  is  a  small  reddish-gray  mass  occupy- 
ing the  sella  turcica  of  the  sphenoid  bone.  It  consists  of  the  pituitary  gland 


THE    PITUITARY   BODY 


495 


proper  or  anterior  lobe,  the  pars  anterior,  the  posterior  lobe,  pars  nervosa, 
and  an  intermediate  epithelial  zone  known  as  the  pars  intermedia. 

The  pars  anterior  or  glandular  lobe  of  the  pituitary  and  also  the  pars 
intermedia  are  derived  from  tissue  that  has  its  embryological  origin  from 
the  epithelial  pouch  of  the  roof  of  the  buco-pharyngeal  cavity.  The  pars 
intermedia  differentiates  out  of  this  common  mass  and  comes  to  invest  the 
infundibular  portion,  apparently  sending  cells  into  its  trabeculae.  Three 
types  of  cells  have  been  described  for  the  pars  amerior;  i,  neutrophiles,  2, 
acidophiles,  and  3,  basophiles.  The  pars  nervosa  consists  of  a  loose  neuroglia 
like  network  arising  out  of  the  base  of  the  third  ventricle.  The  net  exhibits 
open  spaces  more  or  less  filled  with  ingrowths  of  pars  intermedia  cells. 
The  pars  intermedia  (see  Fig.  319)  shows  cavities  filled  with  colloid  being 


FIG.  318. — Mesial  sagitta1  section  of  the  pituitary.     B,  third  ventricle.     AL  and  PL. 
anterior  and  posterior  lobes  respectively.     (Gushing  and  Goetsch.) 

discharged  into  the  spaces  of  the  pars  nervosa.  It  is  thought  that  this 
colloid  may  be  directly  or  indirectly  the  source  of  the  active  hormones  of 
the  posterior  division  of  the  pituitary. 

a.  The  Function  of  the  Pituitary. — The  whole  problem  of  the  function  of 
the  pituitary  body  has  been  extensively  studied  physiologically  and  sur- 
gically by  Gushing.  This  author  has  obtained  concordant  results  by  his 
operative  study  on  dogs,  and  by  the  operative  and  clinical  treatment  of  a 
long  series  of  human  patients.  That  the  pituitary  is  absolutely  necessary 
to  life  is  now  recognized.  A  number  of  functional  disturbances  have  been 
traced  to  abnormal  states  of  this  organ,  and  evidence  is  accumulating  to 


METABOLISM,    NUTRITION,    AND   DIET 

show  that  the  anterior  and  posterior  lobes  are  not  concerned  with  identical 
functions.  Attempts  to  arrive  at  the  knowledge  of  the  proper  physiology 
of  the  organ  have  been  made  primarily  along  three  experimental  lines;  i, 
Removal  of  the  gland  in  whole  or  in  part;  2,  Transplantation;  3,  The  feed- 
ing or  the  injection  of  extracts. 


FIG.  319. — Enlarged  view  of  the  square  marked  on  Fig.  318.     (Gushing  and  Goetsch.) 

b.  Functional  Disturbances  After  Complete  Removal  of  the  Pituitary. — 
Removal  of  the  gland  is  followed  by  death  of  the  operated  animals,  in  old 
dogs  in  from  three  to  five  days,  but  puppies  survive  ten  to  thirty  days 
(Gushing).  The  acute  symptoms  are:  insensitiveness,  slow  pulse  and 
respiration,  fall  in  body  temperature,  apathy,  coma,  and  death. 

If  a  small  part  of  the  gland  is  left  so  that  only  a  condition  of  hypopitui- 
tarism  is  induced  then  the  symptoms  are  more  pronounced  and  chronic. 


THE    PITUITARY   BODY 


497 


These  are  summarized  by  Gushing:  " Widespread  adiposity,  nutritional 
changes  in  the  skin  and  its  appendages,  disturbance  of  carbohydrate 
metabolism,  of  body  temperature,  of  growth  and  of  renal  secretion. 
Sexual  inactivity  or  actual  atrophy  of  the  reproductive  glands  was 
observed;  and,  indeed,  modification  of  most  of  the  other  ductless  glands 
proved  to  be  histologically  demonstrable."  In  puppies  there  was  per- 
sistent "sexual  infantilism,"  " skeletal  undergrowth"  and  "psychic  dis- 
orders." These  experimental  results  throw  light  on  a  great  mass  of 
clinical  evidence  which  Gushing  has  gathered  and  interpreted. 


FIG.  320. — High  magnification  of  the  square  marked  on  Fig.  319.  C,  Colloidol  ves- 
icles apparently  discharging  from  C  into  the  tissue  spaces  of  the  pars  nervosa.  H,  Hyaline 
bodies.  (Gushing  and  Goetsch). 

c.  Action  of  Pituitary  Grafts,  Extracts,  Etc. — Transplantation  of  the  pitui- 
tary into  the  cerebral  cortex  prolonged  the  life  of  Cushing's  animals.  The 
gland  gave  histological  evidence  of  viability  for  a  month.  Grafts  have  not, 
however,  been  umformally  and  permanently  beneficial,  and  do  not  produce 
evidences  of  hyperpituitarism. 

32 


498  METABOLISM,    NUTRITION,    AND    DIET 

In  1911  Gushing  for  the  first  time  transplanted  the  pituitary  of  a  child 
into  an  adult  man  suffering  from  lack  of  pituitary  secretion.  A  pituitary 
cyst  had  been  removed.  The  patient  was  temporarily  but  completely  aroused 
from  the  "profound  torpor"  of  hypopituitarism  to  a  temporary  state  of  normal 
mental  and  physical  vigor  by  injections  of  gland  extracts.  After  the  gland 
was  grafted  the  extracts  were  ceased  and  the  man  remained  in  his  recovered 
state  of  health. 

In  addition  to  the  general  nutritional  improvement  from  the  action  of 
extracts  it  is  now  known  that  smooth  muscular  tissues  are  stimulated,  in 
this  case  by  posterior  lobe  extracts.  Uterine  contractions  in  time  of  atonia 
in  particular,  are  sharply  stimulated,  pituitrin  producing  this  action  strongly. 
The  reaction  in  the  blood  vessels  leads  to  rise  of  blood-pressure,  increase  of 
vagus  tone,  and  augmentation  of  the  heart  beat.  Pituitrin  and  posterior 
lobe  extracts  are  vigorous  galactogogues. 

The  feeding  of  extracts  has  been  less  positive,  though  prolonged  feeding 
has  seemed  to  aid  in  preventing  the  symptoms  of  hypopituitarism  in 
alleviating  obesity  in  particular. 

e.  Acromegaly  or  Hyperpituitarism. — Hypertrophy  of  the  pituitary  leads 
to  a  great  acceleration  in  skeletal  growth  or  gigantism.  This  we  have  not 
been  able  to  prove  by  experimental  means  but  must  now  assume  to  be 
established  by  clinical  and  pathological  findings.  The  accelerated  skeletal 
growth  may  occur  either  in  childhood,  youth  or  after  full  development,  for 
example,  Cushing's  cases  numbers  XXXII  and  I,  which  began  to  grow 
rapidly  at  fifteen  and  twenty-five  years  of  age  respectively. 

Hyperpituitarism  with  its  giant  skeletal  growth  shows  a  tendency  to  pass 
over  to  hypopituitarism  with  its  attendant  nutritional  disorders  of  obesity, 
carbohydrate  tolerance,  and  disturbance  of  correlated  internal  secreting 
glands  and  sex  gonads. 

Acromegaly  as  such  seems  due  to  a  disturbance  of  the  pars  anterior,  while 
the  other  nutritional  derangements  are  attributed  to  disturbance  of  the  pars 
intermedia  and  pars  nervosa. 

The  Reproductive  Glands. — The  ovary  and  the  testis  are  undoubtedly 
concerned  with  metabolism  in  the  body.  Numerous  observations,  not  only 
of  the  sex  gonads  but  of  other  parts  of  the  reproductive  system,  show  an  inti- 
mate relation  of  this  apparatus  to  the  health  and  normal  activities  of  other 
parts  of  the  body. 

a.  The  Testis. — It  has  been  shown  repeatedly  that  extracts  of  the  tissue 
of  the  testis  fed,  or  hypodermically  injected  when  purified,  increased  the 
vigor  both  of  the  muscular  and  of  the  nervous  systems.  Ergograms  show  an 
increase  in  muscular  power.  Spermin  isolated  from  the  testis  is  claimed  by  its 
discoverer  to  produce  these  beneficial  effects.  Sterilization  by  the  removal  of 
the  testis  in  domestic  animals  is  followed  by  an  entire  change  in  the  physical 
development  of  the  animal,  especially  in  the  so-called  secondary  sexual 


THE    REPRODUCTIVE    GLANDS  499 

characters.  Such  animals  show  less  vigor  and  less  muscular  power.  Sterili- 
zation of  the  male  without  removal  of  the  gonads  is  not  followed  by  the  usual 
loss  of  vigor  and  of  the  typical  male  form  and  characteristics.  This  is  ascribed 
to  the  preservation  of  the  testis  which  is  thought  to  produce  internal  secretion 
reacting  through  the  nutritional  processes  of  the  body. 

b.  The  Ovaries. — The  ovaries  not  only  possesses  the  germ  cells,  the  ova, 
but  their  structure  is  characterized  by  other  types  of  cells — the  interstitial, 
stroma,  and  f ollicular  cells.  When  a  graafian  follicle  ruptures  and  discharges 
its  ovum,  cells  rapidly  grow  and  fill  the  cavity  forming  the  corpus  luteum. 

The  surgical  removal  of  the  ovaries  results  in  disturbance  of  the  periodic 
menstrual  phenomenon,  not  because  of  the  loss  of  the  ova,  but  because  the 
secretin  fails  which  maintains  the  normal  periodic  reactions  of  the  uterus. 
There  are  also  in  the  human  disturbances  in  nutrition  from  this  cause,  i.e., 
a  decrease  in  oxidations,  tendency  toward  obesity,  marked  nervous  symptoms, 


.  FIG.  321. — The  galactagogic  action  of  involuting  uterus  of  the  cat  after  intravenous 

injection  of  saline  extracts.     First  tracing  from  above,  blood-pressure;  second,  drops  of 

ulk  from  a  hollow  needle  inserted  into  a  mamma;  third,  drops  from  a  wick  inserted  into 

two  mammas  that  have  been  split  open  to  remove  the  effect  of  the  contractions  of  the  muscles 

of  the  gland.     (Mackenzie.) 

etc. ,  These  symtoms  are  reduced  or  entirely  disappear  on  grafting  a  portion 
of  the  gland,  and  the  disturbed  menstruation  becomes  regular  again.  Human 
ovaries  have  been  transplanted  to  patients  after  previous  ovariotomy  with  a 
recovery  of  normal  body  functions  and  improvements  in  health.  Experi- 
ments by  Lowey  and  Richter  indicate  that  oxidations  in  the  body  are  greatly 
increased  on  feeding  ovarian  extract  to  ovariotomized  animals.  Removal 
of  the  ovaries  of  hens  leads  to  a  loss  of  feminine  characters,  i.e.,  the  develop- 
ment of  spurs,  cock-like  comb  and  wattles,  hackle,  etc.  Castrated  female 
ducks  are  also  said  to  develop  male-like  characters.  Guthrie  suggests  that 
this  may  be  "  an  unmasking  of  characters,"  since  we  do  not  know  the  asexual 
type  in  the  higher  animals  by  any  positive  evidence. 


5OO  METABOLISM,   NUTRITION,    AND    DIET 

c.  Corpus  Luteum. — The  cells  of  the  corpus  luteum  are  now  believed  to 
produce  an  internal  secretion  or  hormone.  This  body  which  develops 
rapidly  after  the  rupture  of  the  graafian  follicles  ordinarily  persists  only  two 
or  three  weeks.  When,  however,  the  ovum  becomes  fertilized  and  attaches 
to  the  uterine  wall  and  its  development  proceeds,  then  the  corpus  luteum 
enlarges  and  persists  throughout  the  gestation  period.  Extracts  of  this 
body  were  found  by  Mackenzie  to  be  galactagogues. 

The  Phenomena  of  Gestation  and  Lactation  in  Relation  to  Internal 
Secretions. — Physiologists  are  accumulating  more  and  more  evidence  sup- 
porting the  view  that  hormones  exert  a  strong  influence  in  the  control  of 
the  process  of  gestation  and  lactation. 

During  gestation,  aside  from  the  mere  fact  of  the  development  of  the 
embryo,  there  are  the  following  special  conditions  to  be  given  considera- 
tion, namely,  the  persistence  of  the  corpus  luteum,  the  great  enlargement  of 
the  uterine  mass,  the  development  of  the  mammary  glands,  to  which  may  be 
added  for  this  consideration,  the  development  of  the  placenta  and  of  the  fetus 
itself.  During  this  period  there  is  a  profound  disturbance  of  many  nutri- 
tional factors  especially  affecting  the  correlations  of  the  digestive,  the  muscular, 
and  the  nervous  systems. 

After  parturition  the  following  changes  in  the  picture  occur,  namely,  the 
discharge  of  the  placenta  which  implies  retrogression  in  this  organ,  involuting 
changes  in  the  uterus,  and  active  lactation.  The  latter  phenomenon  is  of 
special  physiological  interest.  The  growth  of  the  gland  without  lactation 
during  gestation  and  the  abrupt  development  of  the  phenomenon  of  lactation 
apparently  depends  strictly  upon  the  hormones.  As  will  be  seen  later,  the 
mammary  gland  tissue  is  poorly  supplied  with  nerves,  apparently  having  no 
secretory  nerves.  It  would  seem  that  we  have  here  in  the  first  period  a 
group  of  hormones  stimulating  growth,  possibly  associated  with  inhibitive 
hormones  for  milk  secretion.  After  gestation  active  galactagogue  hormones 
control  the  phenomenon.  Such  hormones  are  known  to  come  from  the 
involuting  uterus,  corpus  luteum,  posterior  lobe  of  the  pituitary,  and  from 
the  mammary  gland  itself.  The  present  view  is  that  an  inhibitive  influence  is 
exerted  by  the  hormones  from  the  placenta  and  the  developing  fetus  on 
milk  production  during  the  gland  growth.  Our  information  concerning 
this  complicated  question  is,  however,  at  the  present  time  inadequate. 

There  are  other  organs  whose  function  is  still  obscure  but  in  which 
the  indirect  evidence  points  to  an  influence  on  metabolism  at  one  stage  or 
another  of  the  existence  of  the  animal  body.  Enough  has  been  given  here 
to  show  that  the  interrelation  of  the  organs  of  the  body  is  extremely  complex 
in  so  far  as  metabolism  is  concerned.  It  is  not  enough  simply  to  know  the 
foods  and  their  composition.  The  whole  complex  of  intermediary  metabol- 
isms and  metabolites  and  their  influences  must  constantly  be  taken  into 
consideration. 


CHAPTER  XII. 
ANIMAL  HEAT. 

HEAT  is  produced  by  the  metabolism  of  the  tissues  of  the  body.  In  man 
and  in  such  animals  as  are  called  warm-blooded,  i.e.,  only  mammals  and 
birds,  there  is  an  average  body  temperature  which  is  maintained  with  only 
slight  variations  in  spite  of  changes  in  their  environment.  The  possible 
variations  above  and  below  this  average  are  comparatively  slight.  The 
average  temperature  in  all  mammals  and  birds  is  not  the  same,  for,  as  we 
shall  see,  the  average  temperature  of  man  is  37°  C.  (98.6°  F.),  in  some  birds 
it  is  as  high  as  44°  C.,  while  in  the  wolf  it  is  said  to  be  under  36°  C. 

The  average  temperature  of  the  human  body  in  those  internal  parts  which 
are  most  easily  accessible,  as  the  mouth  and  rectum,  is  from  36 . 9°  to  37 . 4°  C. 
(98 . 5°  to  99 . 5°  F.).  In  different  parts  of  the  external  surface  of  the  human 
body  the  temperature  varies  only  to  the  extent  of  one  or  two  degrees  centi- 
grade, when  all  are  alike  protected  from  cooling  influences;  and  the  differ- 
ence which  under  these  circumstances  exists  depends  chiefly  upon  the 
different  degrees  of  blood  supply.  In  the  axilla  and  in  the  groin,  the  most 
convenient  situations,  under  ordinary  circumstances,  for  examination  by  the 
thermometer,  the  average  temperature  is  37°  C.  (98.6°  F).  In  different 
internal  parts,  the  variation  is  one  or  two  degrees;  those  parts  and  organs 
being  warmest  which  contain  most  blood,  and  in  which  there  occurs  the 
greatest  amount  of  chemical  change,  e.g.,  the  muscles  and  the  glands.  The 
temperature  is  highest  when  they  are  in  a  condition  of  activity.  Those  tis- 
sues which  subserve  only  a  mechanical  function  and  are  the  seat  of  least  ac- 
tive circulation  and  chemical  change  are  the  coolest.  These  differences  of 
temperature,  however,  are  actually  but  slight,  on  account  of  the  provisions 
which  exist  for  maintaining  uniformity  of  temperature  in  different  parts. 

The  average  temperature  of  a  healthy  body  varies  somewhat  according 
to  age,  sex,  time  of  day,  climate,  etc.  The  mean  temperature  is  said  to  be 
slightly  higher,  0.5°  C.,  in  young  children  and  in  old  persons  than  in  adults. 
It  is  perhaps  very  slightly  higher  in  women  than  in  men,  in  warm  climates 
than  in  cold,  in  winter  than  in  summer.  It  varies  slightly  at  different  times 
in  the  day,  especially  during  sleep  when  metabolism  is  at  a  low  ebb. 

Diurnal  Temperature  Variations. — The  discussion  presented  in  this 
paragraph  is  abstracted  from  an  excellent  paper  on  the  subject  by  Professor 
Gibson.  "Certain  features  of  daily  rhythm  are  generally  recognized, 
such  as  the  rise  of  temperature  during  the  forenoon  and  afternoon,  and 

501 


502  ANIMAL   HEAT 

the  fall  during  the  evening  and  early  morning  hours.  A  study  of  the 
literature  indicates  considerable  differences  in  the  incidence  of  the  maxi- 
mum and  minimum  temperature,  depending  undoubtedly  upon  a  variety 
of  causes.  Despite  differences  as  regards  dietetic  habits,  work,  age, 
etc.,  the  time  of  maximum  temperature  may  be  limited  broadly  between 
4  and  8  P.M.,  and  that  of  the  minimum  between  2  A.M.  and  7  A.  M.,  with  an 
average  range  of  variation  of  over  i°  C.  in  rectal  observations.  While  a 
hot  or  cold  climate  may  determine  minimal  variations  in  the  peripheral  tem- 
perature of  the  body,  the  internal  temperature,  as  measured  in  the  rectum, 
appears  to  suffer  only  very  slight  modifications  from  such  causes.  The  rise 
in  temperature  incidental  to  muscular  exertion  is  transitory.  The  follow- 
ing quotations  serve  to  indicate  prevalent  views  regarding  the  diurnal 
rhythm.  Pembrey  writes:  'As  regards  the  causes  of  the  daily  varia- 
tion in  temperature,  muscular  activity  and  food  appear  to  be  the  most  impor- 
tant factors.  In  ordinary  life  man  is  most  active  and  takes  food  during  the 
day,  and  is  least  active  during  the  night.  .  .  .  We  may  conclude  that  the 
daily  variation  in  temperature  is  one  of  the  features  of  a  corresponding  varia- 
tion in  the  activity  of  the  tissues  of  the  body,  as  shown  by  the  rate  of  the  con- 
traction of  the  heart,  the  frequency  of  respiration,  the  intake  of  oxygen,  the 
output  of  carbon  dioxide,  the  discharge  of  urea,  and  the  capacity  for  muscu- 
lar work.'  Jiirgensen's  observations  on  fasting  subjects  in  which  a  more 
or  less  "normal"  curve  of  temperature  variations  was  maintained  exclude 
the  diet  factor  from  a  preponderating  role.  In  estimating  the  relative  im- 
portance of  the  bodily  activities  as  distinguished  from  other  environmental 
conditions,  studies  on  the  influence  of  the  inversion  of  the  daily  routine  have 
been  made.  Those  of  Benedict  claim  special  interest,  because  the  observa- 
tions were  practically  continuous  over  long  periods  of  time.  Such  an  ar- 
rangement was  made  possible  by  the  use  of  a  specially  devised  electric 
resistance  thermometer  reading  to  0.01°  C.,  which  can  be  inserted  10  cm. 
to  15  cm.  in  the  rectum  and  retained  there  without  inconvenience  during 
both  waking  and  sleeping  hours.  Two  subjects  were  carefully  observed  by 
Benedict:  one  a  person  usually  working  during  the  day,  but  made  to  work 
at  night  and  sleep  during  the  day  for  a  series  of  consecutive  days;  the  other, 
an  individual  long  accustomed  to  night-work  in  the  capacity  of  a  night 
watchman.  From  the  results  of  these  observations  it  has  been  assumed  that 
the  general  form  of  the  night  curve  remains  practically  intact,  even  when 
the  daily  routine  is  inverted,  indicating  a  fixity  of  rhythm  that  is  difficult  to 
explain.  As  Benedict  says:  'Why  the  temperature  of  the  human  body 
reaches  a  minimum  at  2  A.M.  to  6  A.M.,  independent  of  whether  the  subject  is 
sleeping  soundly  in  the  recumbent  position  or  whether  he  is  awake  and 
sitting,  or  even  standing  and  walking,  is  a  problem  that  calls  for  extended 
research.'  However,  the  transposition  of  the  daily  routine  through  a  period 
of  practically  half  a  day,  experienced  as  the  result  of  the  time  changes 


VARIATION    IN    THE    LOSS    OF    HEAT  503 

during  the  trip  from  New  Haven  to  Manila,  produces  a  corresponding  and 
coincident  shifting  of  the  rhythmic  temperature  changes,  so  that  the  normal 
character  of  the  variations  is  always  preserved." 

Heat-producing  Organs. — Heat  is  liberated  in  the  body  wherever 
oxidative  metabolism  takes  place.  Of  all  the  tissues  of  the  body  muscular 
tissue  is  conspicuous  for  its  mass  and  for  its  activity.  It  is  evidently  the  great 
heat-producing  tissue.  The  manifestation  of  muscular  energy  is  always  ac- 
companied by  the  evolution  of  heat  and  the  production  of  carbon  dioxide. 
This  production  of  carbon  dioxide  goes  on  while  the  muscles  are  in  mechani- 
cal rest,  only  in  a  less  degree  than  that  which  is  noticed  during  muscular 
activity,  and  so  it  is  certain  that  an  active  katabolism  is  going  on  in  resting  as 
well  as  in  contracting  muscles.  This  katabolism  is  a  source  of  much  heat,  and 
so  the  total  amount  of  heat  produced  in  the  muscular  tissues  per  day  must 
be  very  great.  It  has  been  calculated  that,  even  neglecting  the  heat  produced 
by  the  quiet  katabolism  of  muscular  tissue,  the  amount  of  heat  generated  by 
muscular  activity  would  supply  the  principal  part  of  the  total  heat  produced 
within  the  body.  The  heart,  as  a  special  muscle,  deserves  particular  mention 
since  it  is  in  constant  vigorous  activity.  All  its  energy  is  ultimately  converted 
into  heat,  accounting  for  about  5  per  cent,  of  the  total  heat  of  the  body.  The 
secreting  glands,  and  principally  the  liver  as  being  the  largest  and  most  ac- 
tive, come  next  to  the  muscles  and  heart  as  heat-producing  tissues.  It  has 
been  found  by  experiment  that  the  blood  leaving  the  glands  is  considerably 
warmer  than  that  entering  them.  The  metabolism  in  the  glands  is  very 
active;  and  the  more  active  the  katabolism,  the  greater  the  heat  produced. 

It  must  be  remembered,  however,  that  although  the  organs  mentioned  are 
the  chief  heat-producing  parts  of  the  body,  all  living  tissues  contribute  their 
quota,  and  this  in  direct  proportion  to  their  activity.  The  blood  itself  is  also 
the  seat  of  katabolism  and,  therefore,  of  the  production  of  heat;  but  the 
share  which  it  takes  in  this  respect,  apart  from  the  tissues  in  which  it  circu- 
lates, is  very  inconsiderable. 

Regulation  of  the  Temperature  of  the  Human  Body. — The  average 
temperature  of  the  body  is  maintained  under  different  conditions  of  external 
circumstance  by  mechanisms  which  permit  of  (i)  variation  in  the  loss  of  heat, 
and  (2)  variations  in  the  production  of  heat.  In  healthy  warm-blooded  ani- 
mals the  loss  and  gain  of  heat  are  so  nearly  balanced  one  by  the  other  that, 
under  all  ordinary  circumstances,  a  uniform  temperature,  within  a  degree  or 
two,  is  preserved. 

Variation  in  the  Loss  of  Heat. — The  loss  of  heat  from  the  human 
body  is  principally  regulated  by  the  amount  given  off  by  radiation  and  con- 
duction from  its  surface,  by  means  of  the  constant  evaporation  of  water  from 
the  same  part,  heat  being  thus  rendered  latent,  and  to  a  much  less  degree  by 
loss  from  the  air-passages.  In  each  act  of  respiration,  heat  is  lost  to  a  greater 
or  less  extent  according  to  the  temperature  of  the  atmosphere;  unless  indeed 


504  ANIMAL  HEAT 

the  temperature  of  the  surrounding  air  exceeds  that  of  the  blood.  We  must 
remember,  too,  that  all  food  and  drink  which  enter  the  body  at  a  lower  tem- 
perature abstract  a  small  measure  of  heat;  while  the  urine  and  feces  which 
leave  the  body  at  about  its  own  temperature  are  also  means  by  which  a  certain 
small  amount  of  heat  is  lost. 

Heat  Lost  from  the  Surface  of  the  Body. — By  far  the  most  impor- 
tant loss  of  heat  from  the  body,  probably  90  per  cent,  and  upward  of  the 
whole  amount,  is  that  which  takes  place  by  radiation,  conduction,  and  the 
evaporation  of  moisture  from  the  skin.  The  actual  figures  for  ordinary 
conditions  are  as  follows:  For  every  100  calories  of  heat  produced,  2.6  are 
lost  in  heating  the  food  and  drink;  2.6  in  heating  the  air  inspired;  14.7  in 
evaporation;  and  80.  i  by  radiation  and  conduction.  During  increased 
activity  of  the  body  the  proportion  of  heat  lost  by  evaporation  is  greatly 
increased.  The  means  by  which  the  skin  is  able  to  act  as  one  of  the  most 
important  organs  for  regulating  the  temperature  of  the  blood  are,  i,  that 
it  offers  a  large  surface  for  radiation,  conduction,  and  evaporation;  2,  that  it 
contains  a  large  but  adjustable  amount  of  blood,  and  the  quantity  of  blood 
is  greater  under  those  circumstances  which  demand  a  loss  of  heat  from  the 
body,  and  vice  versa,  which  gives  a  means  for  varying  the  loss  of  heat  by 
radiation  and  conduction;  3,  that  it  contains  the  sweat  glands,  which  dis- 
charge a  quantity  of  moisture  to  be  evaporated  from  its  surface. 

The  circumstance  which  directly  determines  the  quantity  of  blood  in  the 
skin  is  that  which  governs  the  supply  of  blood  to  all  the  tissues  and  organs 
of  the  body,  namely,  the  power  of  the  vaso-motor  nerves  to  cause  a  greater 
or  less  tension  of  the  muscular  element  in  the  walls  of  the  arteries,  and,  in 
correspondence  with  this,  a  lessening  or  increase  of  the  caliber  of  the  vessel, 
accompanied  by  a  less  or  greater  current  of  blood.  A  warm  or  hot  atmos- 
phere so  acts  on  the  sensory  nerves  of  the  skin  as  to  lead  to  a  reflex  relaxa- 
tion of  the  muscular  fiber  of  the  blood  vessels;  as  a  result,  the  skin  becomes 
full-blooded,  relatively  hot,  and  moist  from  sweating;  and  much  heat  is  lost. 
With  a  low  temperature  the  blood  vessels  shrink,  and  with  the  consequently 
diminished  blood  supply,  the  skin  becomes  pale,  cold,  and  dry,  an  effect 
produced  through  the  vascular  centers  in  the  medulla  and  spinal  cord. 

The  activity  of  the  sweat  glands  of  the  skin  is  also  regulated  reflexly 
through  the  sweat  centers.  The  increased  blood  supply  just  described  is 
favorable  to  increased  production  of  sweat  by  the  sweat  glands.  Thus, 
by  means  of  the  self-regulation  the  skin  becomes  the  most  important  of  the 
means  by  which  the  temperature  of  the  body  is  regulated. 

The  relative  loss  of  heat  by  the  means  given,  i.e.,  radiation,  conduction, 
and  evaporation,  will  depend  on  two  factors:  first,  the  relative  temperature 
of  the  body  to  the  surrounding  air;  and,  second,  the  humidity  of  the  air.  If 
the  atmospheric  temperature  is  the  same  as  that  of  the  body,  of  course  there 
will  be  no  loss  of  heat  by  radiation  and  convection;  if  the  air  temperature  is 


HEAT    LOST    FROM    THE    SURFACE    OF    THE    BODY  505 

higher,  there  will  be  an  actual  gain.  When  the  humidity  of  the  air  is  great, 
there  will  be  reduced  evaporation  of  perspiration  and  consequent  diminished 
heat  loss.  If  we  assume  a  moisture-saturated  air  at  the  body  temperature, 
then  heat  loss  becomes  impossible  and  the  temperature  of  the  body  will 
rise.  This  is  why  a  hot  moist  climate  is  so  oppressive,  while  a  hot  but  dry 
atmosphere  is  readily  borne  by  the  human  body.  The  amount  of  heat 
required  to  evaporate  i  c.c.  of  water  is  536  small  calories,  hence  an  increase 
in  the  evaporation  of  perspiration  readily  compensates  for  a  decrease  in  the 
loss  of  heat  by  radiation  and  convection. 

Many  examples  may  be  given  of  the  power  which  the  body  possesses  of 
resisting  the  effects  of  a  high  temperature,  in  virtue  of  evaporation  from  the  skin. 
Blagden  and  others  supported  a  temperature  varying  between  92°  to  100°  C. 
(i98°-2i2°F.)  in  dry  air  for  several  minutes;  and  in  a  subsequent  experiment 
he  remained  eight  minutes  in  a  temperature  of  126.5°  C.  (260°  F.).  "The 
workmen  of  Sir  F.  Chantrey  were  accustomed  to  enter  a  furnace,  in  which  his 
molds  were  dried,  while  the  floor  was  red-hot,  and  a  thermometer  in  the 
air  stood  at  177  .8°  C.  (350°  F.),  and  Chabert,  the  fire  king,  was  in  the  habit  of 
entering  an  oven  the  temperature  of  which  was  from  205°— 315°  C.  (400°— 
600°  F.)."  (Carpenter.) 

But  such  heats  are  not  tolerable  when  the  air  is  moist  as  well  as  hot,  so  as 
to  prevent  evaporation  from  the  body.  C.  James  states  that  in  the  vapor 
baths  of  Nero  he  was  almost  suffocated  in  a  temperature  of  44.5°  C.  (112°  F.), 
while  in  the  caves  of  Testaccio,  in  which  the  air  is  dry,  he  was  but  little  incom- 
moded by  a  temperature  of  80°  C.  (176°  F.).  In  the  former,  evaporation  from 
the  skin  was  impossible;  in  the  latter  it  was  abundant,  and  the  layer  of  vapor 
which  would  rise  from  all  the  surface  of  the  body  would,  by  its  very  slowly 
conducting  power,  defend  it  for  a  time  from  the  full  action  of  the  external  heat. 

Man  is  able  by  suitable  clothing  to  increase  or  to  diminish  the  amount  of 
heat  lost  by  the  skin.  There  are  baths  and  other  means  which  man  and 
animals  instinctively  adopt  for  lowering  the  temperature  when  necessary. 

Although  under  any  ordinary  circumstances  the  external  application  of 
cold  only  temporarily  depresses  the  normal  temperature  to  a  slight  extent,  it 
is  otherwise  in  cases  of  high  temperature  in  fever.  In  these  cases  a  cool 
bath  may  reduce  the  temperature  several  degrees,  and  the  effect  so  pro- 
duced lasts  in  some  cases  for  many  hours. 

Extreme  heat  and  cold  produce  effects  too  powerful,  either  in  raising  or 
lowering  the  heat  of  the  body,  to  be  controlled  by  the  proper  regulating  ap- 
paratus. Walther  found  that  rabbits  and  dogs  kept  exposed  to  a  hot  sun, 
reached  a  temperature  of  46°  C.  (114.8°  F.),  and  then  died.  Cases  of  sun- 
stroke furnish  us  with  several  examples  in  the  case  of  man;  for  it  would  seem 
that  here  death  ensues  chiefly  or  solely  from  elevation  of  the  temperature. 

The  effect  of  mere  loss  of  bodily  temperature  in  man  is  less  well  known 
than  the  effect  of  heat.  From  experiments  by  Walther  it  appears  that  rab- 
bits can  be  cooled  down  to  8 . 9°  C.  (48°  F.)  before  they  die,  if  artificial  respira- 
tion be  kept  up.  Cooled  down  to  17.8°  C.  (64°  F.),  they  eannot  recover 


506  ANIMAL   HEAT 

unless  both  external  warmth  and  artificial  respiration  be  employed.  Rabbits 
not  cooled  below  25°  C.  (77°  F.)  recover  by  external  warmth  alone. 

Loss  of  Heat  from  the  Lungs. — The  lungs  and  air-passages  are  very 
inferior  to  the  skin  as  a  means  for  lowering  the  temperature.  In  giving 
heat  to  the  air  breathed,  the  lungs  stand  next  to  the  skin  in  importance.  As 
a  regulating  power,  the  inferiority  is  very  marked.  The  air  which  is  ex- 
pelled from  the  lungs  leaves  the  body  at  about  the  temperature  of  the  blood, 
and  is  always  saturated  with  moisture.  No  inverse  proportion,  therefore, 
exists,  as  in  the  case  of  the  skin,  between  the  loss  of  heat  by  radiation  and 
conduction,  on  the  one  hand,  and  by  evaporation,  on  the  other.  The  colder 
the  air  and  the  drier,  for  example,  the  greater  will  be  the  loss  in  all  ways. 
Neither  is  the  quantity  of  blood  which  is  exposed  to  the  cooling  influence  of 
the  air  diminished  or  increased  in  the  lungs,  so  far  as  is  known,  in  accord- 
ance with  any  need  in  relation  to  temperature.  It  is  true  that  by  varying  the 
number  and  depth  of  the  respirations,  the  quantity  of  heat  given  off  by  the 
lungs  may  be  made  to  vary  also  for  a  few  minutes.  But  the  respiratory 
passages,  while  they  must  be  considered  important  means  by  which  heat 
is  lost,  are  altogether  subordinate,  in  the  power  of  actively  regulating  the 
temperature. 

The  loss  of  heat  used  to  warm  foods  is  an  obvious  method  of  expenditure 
of  heat  which  may  be  resorted  to,  especially  in  certain  fevers.  The  loss  of 
heat  by  the  excreta  discharged  from  the  body  at  a  high  temperature  must  be 
of  no  use  as  a  means  of  regulating  the  temperature,  since  the  amount  so  lost 
must  be  capable  of  little  variation. 

Variation  in  the  Production  of  Heat. — It  may  seem  to  have  been 
assumed,  in  the  foregoing  pages,  that  the  only  regulating  apparatus  for  tem- 
perature required  by  the  human  body  is  one  that  shall,  more  or  less,  produce 
a  cooling  effect;  as  if  the  amount  of  heat  produced  were  always,  therefore, 
in  excess  of  that  which  is  required.  Such  an  assumption  would  be  incorrect. 
The  body  has  the  power  of  regulating  the  production  of  heat,  as  well  as  its 
loss. 

The  production  of  heat  in  the  body  is  apparently  different  for  each  ani- 
mal; i.e.,  the  absolute  amount  of  heat  set  free  by  different  animals  in  a  given 
period  varies.  Each  individual  has  his  own  coefficient  of  heat  production. 
From  all  that  has  been  said  on  the  subject  it  will  be  seen  that  the  amount  of 
heat  for  all  practical  purposes  depends  upon  the  metabolism  of  the  tissues  of 
the  body;  everything,  therefore,  which  increases  that  metabolism  will  increase 
the  heat  production;  so,  therefore,  the  absolute  amount  of  heat  produced  by  a 
large  animal,  having  a  larger  amount  of  tissues  in  which  metabolism  may 
go  on,  will  be,  ceteris  paribus,  greater  than  that  of  a  small  animal.  But  the 
activity  of  the  tissue  change  in  a  small  animal  may  be  greater  than  in  a  large 
one,  as  measured  per  kilo  of  body  weight,  and  naturally  no  strict  line  can  be 
drawn  between  the  two. 


INFLUENCE    OF    NERVOUS    SYSTEM    ON   HEAT    PRODUCTION       507 

HEAT  PRODUCED  PER  KILO  PER  HOUR.     (MuNK.) 

Man i .  o  Calories. 

Dog  (large) 1.7  Calories. 

Dog  (small) 3.8  Calories. 

Guinea-pig 7.5  Calories. 

Rat 11.3  Calories. 

Mouse 19.0  Calories. 

Sparrow 35-5  Calories. 

The  ingestion  of  foods  increases  the  metabolism  of  the  tissues.  As  one 
would  expect,  the  rate  of  heat  production  is  found  by  experiment  upon  the 
dog  to  be  increased  after  a  meal,  reaching  its  height  about  six  hours  after  a 
meal. 

It  has  also  been  experimentally  ascertained  that  the  rate  of  heat  produc- 
tion varies  with  the  kind  of  food  taken:  for  example,  if  sugar  be  added  to  the 
meal  of  meat  given  to  the  dog,  the  height  of  maximum  production  is  reached. 
It  is  often  said  that  the  various  nations  have  found  by  experience  what  food 
is  most  suitable  for  the  climate  in  which  they  live,  and  that  such  experience 
can  be  trusted  to  regulate  the  quantity  consumed.  Although  there  have 
been  no  very  conclusive  experiments  to  prove  the  view,  yet  it  is  a  matter  of 
general  observation  that  in  northern  climates  and  in  colder  seasons  the  quan- 
tity of  food  taken  is  greater  than  in  warmer  climates  or  in  warmer  seasons. 
Moreover,  the  kind  of  food  is  different.  For  example,  persons  living  in  the 
colder  climates  require  much  fat  in  order  to  produce  the  requisite  amount  of 
heat. 

Influence  of  the  Nervous  System  on  Heat  Production. — The  influence 
of  the  nervous  system  in  modifying  the  production  of  heat  must  be  very 
important,  as  upon  the  nervous  influence  depends  the  amount  of  the  metab- 
olism of  the  tissues.  The  experiments  and  observations  which  best  illus- 
trate it  are  those  showing,  first,  that,  when  the  supply  of  nerves  to  a  part  is 
cut  off,  the  temperature  of  that  part  falls  below  its  ordinary  degree  after  a 
time;  and,  second,  that  when  there  is  severe  injury  to  or  removal  of  the 
nervous  centers  the  temperature  of  the  body  rapidly  falls,  even  though  arti- 
ficial respiration  be  performed,  the  circulation  maintained,  and  to  all  appear- 
ance the  ordinary  conditions  for  chemical  changes  in  the  body  be  completely 
maintained. 

There  is  a  heat-regulating  nervous  apparatus  closely  comparable  to  that 
which  regulates  the  secretion  of  saliva  or  of  sweat,  by  means  of  which  the  pro- 
duction of  heat  in  the  warm-blooded  animals  is  increased  or  diminished,  as 
occasion  requires.  This  apparatus  probably  consists  of  a  center  or  centers 
in  the  brain  which  may  be  reflexly  stimulated,  as,  for  example,  by  impulses 
from  the  skin,  and  which  act  through  special  nerves  supplied  to  the  various 
tissues.  The  evidence  upon  which  the  existence  of  this  regulating  appara- 
tus is  assumed  is  the  marked  effect  in  the  increase  of  the  oxygen  consumed  by 


508  ANIMAL  HEAT 

a  warm-blooded  animal  when  exposed  to  cold,  and  the  corresponding  increase 
in  the  output  of  carbon  dioxide,  indicating  that  there  is  an  increase  of  the 
metabolism  and  so  an  increased  production  of  heat  under  such  circumstances, 
and  not  a  mere  diminution  of  the  amount  of  heat  lost  by  the  skin,  etc.  A 
cold-blooded  animal  reacts  very  differently  to  exposure  to  cold;  instead  of 
increasing  the  metabolism  as  in  the  case  of  the  warm-blooded  animal,  cold 
diminishes  the  metabolism  of  its  tissues.  It  is  clear,  therefore,  that  in  warm- 
blooded animals  there  is  some  apparatus  not  possessed  by  cold-blooded  ani- 
mals, which  counteracts  the  effects  of  cold.  In  warm-blooded  animals  poi- 
soned by  curara,  or  in  which  section  of  the  medulla  has  been  done,  it  has  been 
found  that  this  regulating  apparatus  is  no  longer  in  action,  and  under  such 
circumstances  no  difference  appears  to  exist  between  such  animals  and  those 
which  are  naturally  cold-blooded.  Warmth  increases  their  temperature, 
and  cold  lowers  it,  and  with  this  there  is,  of  course,  evidence  of  diminished 
metabolism. 

The  explanation  of  these  experiments  is  that  in  such  animals  the  connec- 
tion between  the  skin  and  the  muscles  through  the  nervous  chain,  such  as 
a  thermotaxic  nervous  apparatus  might  be  supposed  to  afford,  is  broken 
either  at  the  termination  of  the  nerves  in  the  muscles  (curara)  or  at  the  sec- 
tioned point  of  the  bulb. 

The  position  of  these  hypothetical  centers  is  a  matter  of  some  difference 
of  opinion.  It  has  been  demonstrated  that  stimulation  of  certain  parts  of 
the  brain  may,  among  other  symptoms,  produce  increased  metabolism  of  the 
tissues  with  increased  output  of  carbon  dioxide  and  a  raised  temperature: 
the  parts  of  which  this  may  be  asserted  are  parts  of  the  corpus  striatum  and 
of  the  optic  thalamus.  The  general  thermogenic  centers  are  probably  closely 
associated  with  the  motor  centers  of  the  cord  and  brain  stem.  The  thermo- 
regulative  centers  are  the  nuclei  in  the  corpus  striatum  and  optic  thalamus. 
Assuming  a  constant  or  tonic  activity  of  the  thermogenic  regulative  centers, 
it  is  easy  to  understand  the  fall  of  temperature  on  their  destruction  or  on  the 
destruction  of  the  nerve  paths  to  the  active  tissues. 

Experimental  observations,  such  as  have  been  made  upon  animals, 
receive  confirmation  from  the  observations  on  patients  who  suffer  from  fever 
or  pyrexia;  in  them  the  temperature  of  the  body  may  be  raised  several  de- 
grees, as  we  have  already  pointed  out.  This  increase  of  temperature 
might,  of  course,  be  due  to  diminished  loss  of  heat  from  the  skin,  but  this, 
although  a  factor,  is  not  the  only  cause.  The  amount  of  oxygen  taken  in 
and  the  amount  of  carbon  dioxide  given  out  are  both  increased.  With 
this  there  must  be  increased  metabolism  of  the  tissues,  and  particularly  of 
the  muscular  tissues,  since  at  the  same  time  the  amount  of  urea  in  the  urine 
is  increased.  Every  one  is  familiar  with  the  rapid  wasting  which  is  such  a 
characteristic  of  high  fever;  it  indicates  not  only  too  rapid  metabolism  of 
the  body,  but  also  insufficient  time  for  the  tissues  to  again  build  themselves 


INFLUENCE    OF    NERVOUS    SYSTEM    ON   HEAT    PRODUCTION       509 

up.  In  certain  fevers,  therefore,  there  may  be  supposed  to  be  some  inter- 
ference with  the  ordinary  cutaneous  reflex  channels  by  which  the  usual 
temperature  sensory  stimuli  in  the  skin  are  prevented  from  producing  the 
reflexes  that  result  in  diminished  production  of  heat  in  the  muscles  and  other 
tissues.  In  consequence  of  this,  and  in  spite  of  the  condition  of  increased 
heat  of  the  body,  both  at  the  surface  and  in  the  deeper  tissues,  the  production 
of  heat  goes  on  at  an  abnormal  rate.  It  is  not  certain  whether  the  patho- 
logical condition  is  one  which  stimulates  the  thermogenic  center  by  means 
of  which  the  metabolism  of  the  tissues  is  increased,  or  whether  the  normal 
reflexes  which  ordinarily  inhibit  the  activity  of  the  center  when  the  tempera- 
ture rises  fail  to  bring  about  their  usual  reactions.  The  first  is  the  probable 
explanation  of  the  high  fevers  of  certain  toxemias. 

Drugs  may  markedly  interfere  with  the  function  of  the  thermogenic 
mechanism.  For  example,  in  anesthesia  for  operative  purposes  the  tem- 
perature of  the  body  falls  below  the  normal  unless  heat  loss  is  prevented. 
If  too  great  loss  occurs  so  that  the  body  temperature  as  measured  in  the 
rectum  (determined  on  cats)  falls  to  25°  C.,  and  lower,  the  heat  regulating 
mechanism  ceases  to  be  operative.  It  does  not  regain  its  function  without 
the  aid  of  artificial  heat.  But  recovery  of  the  thermogenic  function  may 
be  accomplished  by  artificial  aid  even  after  the  rectal  temperature  has  fallen 
to  as  low  as  16°  C. 


CHAPTER  XIII. 
MUSCLE-NERVE  PHYSIOLOGY. 

The  structure  of  muscle,  of  nerve,  and  of  nerve  relations  to  muscle  are 
all  given  in  considerable  detail  in  Chapter  II,  to  which  the  reader  is  referred 

CHEMICAL  COMPOSITION  OF  MUSCLE. 

Muscle  Plasma. — The  principal  substance  which  can  be  extracted 
from  muscle,  when  examined  after  death,  is  the  protein  body,  myosin. 
This  body  appears  to  bear  somewhat  the  same  relation  to  the  living  muscle 
that  fibrin  does  to  the  living  blood,  since  the  coagulation  of  muscle  after 
death  is  due  to  the  formation  of  myosin.  Thus,  if  coagulation  be  delayed 
by  removing  the  muscles  immediately  that  an  animal  is  killed,  and  rapidly 
cooling  them  to  a  temperature  below  o°  C.  before  the  muscles  themselves 
lose  their  irritability,  it  is  possible  to  express  a  viscid  fluid  of  slightly  alkaline 
reaction,  called  muscle  plasma  (Kiihne).  Muscle  plasma,  if  exposed  to 
the  ordinary  temperature  of  the  air,  undergoes  coagulation  much  in  the  same 
way  as  does  blood  plasma  under  similar  circumstances  when  separated 
from  the  blood  corpuscles  at  a  low  temperature.  The  appearances  presented 
by  the  fluid  during  the  process  are  also  very  similar  to  the  phenomena  of 
blood-clotting,  viz.,  first  of  all  an  increased  viscidity  appears  on  the  surface 
of  the  fluid,  and  at  the  sides  of  the  containing  vessel,  which  gradually  extends 
throughout  the  entire  mass,  until  a  fine  transparent  clot  is  obtained.  In  the 
course  of  some  hours  the  clot  begins  to  contract,  and  to  squeeze  out  of  its 
meshes  a  fluid  corresponding  to  blood  serum.  In  the  course  of  coagulation, 
therefore,  muscle  plasma  separates  into  muscle  clot  and  muscle  serum.  The 
muscle  clot  contains  the  substance  myosin.  It  differs  from  fibrin  in  being 
easily  soluble  in  a  10  per  cent,  solution  of  sodium  chloride.  It  is  insoluble 
in  distilled  water,  and  its  solutions  coagulate  on  application  of  heat;  in  short, 
it  is  a  globulin.  During  the  process  of  clotting  the  reaction  of  the  fluid 
becomes  distinctly  acid. 

The  coagulation  of  muscle  plasma  can  be  prevented  not  only  by  cold, 
but  also,  as  Halliburton  has  shown,  by  the  presence  of  neutral  salts  in  certain 
proportions;  for  example,  of  sodium  chloride,  magnesium  sulphate,  or  sodium 
sulphate.  It  will  be  remembered  that  this  is  also  the  case  with  blood  plasma. 
Dilution  of  the  salted  muscle  plasma  will  produce  slow  coagulation. 

It  is  highly  probable  that  the  formation  of  muscle  clot  is  due  to  the 
presence  of  a  ferment,  myosin  ferment. 

510 


THE    PROPERTIES    OF    LIVING    MUSCLE  511 

The  relation  between  the  proteins  of  living  and  dead  muscle  is  represented 
by  Halliburton  as  follows: 

Proteins  of  the  living  muscle. 


Para-myosinogen.  Myosinogen. 

Soluble  myosin. 


Myosin. 
(The  protein  of  the  muscle  clot.) 

About  75  per  cent,  of  the  total  protein  content  of  living  muscle  is  myo- 
sinogen  and  the  remaining  25  per  cent,  is  para-myosinogen.  These  proteins 
may  be  separated  by  subjecting  the  muscle  plasma  to  fractional  coagulation. 
The  para-myosinogen  coagulates  at  47°  C.  and  the  myosinogen  at  56°  C. 
Para-myosinogen  is  a  globulin  since  it  responds  to  the  precipitation  tests  of 
this  group  of  proteins  and  is  insoluble  in  water.  Myosinogen,  on  the  contrary, 
is  not  a  typical  globulin  since  it  is  soluble  in  water,  but  is  a  pseudo- globulin. 

Other  Constituents  of  Muscle.  —  In  addition  to  muscle  ferments 
already  mentioned,  muscle  contains  certain  proteolytic  enzymes,  as  do 
other  tissues,  an  amylolytic  ferment,  a  maltase  and  a  lactic-acid  forming 
enzyme. 

Certain  acids  are  also  present,  particularly  sarco-lactic  acid. 

Of  carbohydrates,  glycogen  and  dextrose  are  present.  Glycogen  is 
present  in  considerable  amount,  especially  in  the  muscles  of  well-nourished 
animals. 

Nitrogenous  crystalline  bodies,  such  as  creatin,  creatinin,  xanthin,  hypo- 
xanthin,  carnin,  guanin,  urea  in  very  small  amount,  uric  acid,  and  inosinic 
and  phospho-carnic  acids,  are  all  found  on  extracting  dead  muscle. 

Chlorides  and  salts  of  potassium,  calcium,  magnesium,  and  iron  are 
present  in  muscle,  the  chief  of  which  is  potassium  phosphate. 

In  extracts  of  muscles,  especially  of  red  muscles,  there  is  a  certain  amount 
of  hemoglobin,  and  also  of  a  pigment  special  to  muscle,  called  by  McMunn 
myo-hematin,  which  has  a  spectrum  quite  distinct  from  hemoglobin,  viz.,  a 
narrow  band  just  before  D,  two  very  narrow  bands  between  D  and  E,  and 
two  other  faint  bands,  near  E  b,  and  between  E  and  F  close  to  F. 

Fats  occur  in  the  connective  tissue  around  and  between  the  muscle 
fibers,  and  lipoids  and  fat  droplets  also  are  present  within  the  individual 
muscle  fibers. 

THE  PROPERTIES  OF  LIVING  MUSCLE. 

Elasticity. — Muscle  has  a  certain  amount  of  elasticity  during  rest. 
It  admits  of  being  considerably  stretched,  but  returns  readily  and  completely 


512  MUSCLE-NERVE    PHYSIOLOGY 

to  its  normal  condition.  In  the  living  body  the  muscles  are  always  stretched 
somewhat  beyond  their  natural  length,  they  are  always  in  a  condition  of 
slight  tension:  an  arrangement  which  enables  the  whole  force  of  the  con- 
traction to  be  utilized  in  approximating  the  points  of  attachment.  If  the  ex- 
tensibility of  a  given  muscle  be  measured  by  adding  to  it  equal  increments 
of  weight,  it  will  be  found  that  the  extension  or  stretching  is  considerable  at 
first,  but  that  the  amount  decreases  with  each  additional  weight.  If  the 
figures  obtained  be  plotted  on  co-ordinate  paper,  a  curve  approaching  a  parab- 
ola is  obtained,  whereas  a  steel  spring  is  perfectly  elastic  and  gives  a  straight 
line.  When  the  weights  are  removed  from  a  stretched  muscle,  one  by  one, 
the  muscle  regains  its  original  length,  though  slowly.  Extreme  fatigue 
greatly  decreases  the  elasticity,  while  an  increase  of  temperature  increases  it. 

Cardiac  muscle  and  smooth  muscle  both  manifest  elasticity  in  the  same 
manner  as  skeletal  muscle.  In  fact,  the  elasticity  of  the  arterioles  is  chiefly 
due  to  the  smooth  muscle  in  their  walls,  a  fact  that  is  of  great  importance  in 
the  adaptability  of  the  circulatory  apparatus.  The  flexibility  of  the  stom- 
ach, the  urinary  bladder,  etc.,  is  traceable  to  the  same  property  of  their 
muscular  walls. 

Contractility  and  Irritability  of  Muscle. — The  property  of  muscular 
tissue  by  which  its  peculiar  functions  are  exercised  is  its  contractility,  which 
is  excited  by  all  kinds  of  stimuli  applied  either  directly  to  the  muscles  or  in- 
directly to  them  through  the  medium  of  their  motor  nerves.  The  property 
of  the  muscle  which  enables  it  to  respond  to  a  stimulus  is  called  its  irritability. 
This  property,  although  commonly  brought  into  action  through  the  nervous 
system,  is  inherent  in  the  muscular  tissue.  This  is  proven:  i.  By  the  fact 
that  contractility  is  manifested  in  a  muscle  which  is  isolated  from  the  influ- 
ence of  the  nervous  system  by  division  of  the  nerves  supplying  it  so  long  as 
the  natural  tissue  of  the  muscle  is  duly  nourished.  2.  It  is  manifested  in  a 
portion  of  muscular  fiber  in  which,  under  the  microscope,  no  nerve  fiber  can 
be  traced.  3.  Substances  such  as  curara,  which  paralyze  the  nerve  endings 
in  muscles,  do  not  at  all  diminish  the  irritability  of  the  muscle  itself.  4. 
When  a  muscle  is  fatigued,  a  local  stimulation  is  followed  by  a  contraction 
of  a  small  part  of  the  fiber  in  the  immediate  vicinity,  without  any  regard  to 
the  distribution  of  nerve  fibers. 

Forms  of  Stimuli  for  Muscle  or  Nerve. — The  power  of  contraction 
in  voluntary  muscles  is  normally  called  forth  in  the  body  by  nerve  impulses 
which  reach  the  muscles  over  the  motor  nerves.  But  a  muscle  will  respond 
to  stimuli  of  various  kinds,  and  these  stimuli  may  be  applied  directly  to  the 
muscle  or  indirectly  to  the  nerve  supplying  it.  There  are  distinct  advantages, 
however,  in  applying  the  stimulus  to  the  nerve,  as  it  is  more  convenient  as 
well  as  more  potent.  The  stimuli  which  will  produce  contraction  in  a 
muscle  are: 

i.  Mechanical  Stimuli. — A  blow,  pinch,  prick  of  the  muscle  or  its  nerve 


THE    SIMPLE    MUSCLE    CONTRACTION  513 

will  produce  a  contraction,  repeated  on  the  repetition  of  the  stimulus.  If 
applied  to  the  same  point  for  a  number  of  times  such  stimuli  will  soon  destroy 
the  irritability  of  the  preparation. 

2.  Thermal  Stimuli. — If  a  needle  or  glass  rod  be  heated  and  applied  to  a 
muscle  or  its  nerve,  the  muscle  will  contract.     A  temperature  of  over  45°  C. 
will  cause  the  muscles  of  a  frog  to  pass  into  a  condition  known  as  heat  rigor. 
The  sudden  change  of  temperature  acts  as  a  stimulus. 

3.  Chemical  Stimuli. — A  great  variety  of  chemical  substances  will  excite 
the  contraction  of  muscles,  some  substances  being  more  potent  in  irritating 
the  muscle  itself  and  other  substances  having  more  effect  upon  the  nerve. 
Of  the  former  may  be  mentioned  dilute  acids,  salts  of  certain  metals,  e.g., 
zinc,  copper,  and  iron;  to  the  latter  belong  strong  glycerin,  strong  acids, 
ammonia,  bile  salts  in  strong  solution,  etc. 

4.  Electrical  Stimuli. — Any  form  of  electrical  current  may  be  employed 
to  stimulate  a  muscle  to  contract,  but  either  galvanism  or  the  induced  current 
is  usually  chosen.     For  experimental  purposes  electrical  stimuli  are  most 
frequently  used,  as  the  strength  of  the  stimulus  maybe  conveniently  regulated. 
In  order  that  a  stimulus  shall  be  effective,  it  must  have  a  certain  amount  of 
energy  and  the  application  to  the  muscle  must  have  a  certain  abruptness. 
For  example,  a  comparatively  weak  galvanic  current  suffices  to  stimulate  a 
muscle  to  action  when  suddenly  applied  in  full  force.     But  if  the  electric 
current  be  applied  very  gradually,  a  current  many  times  stronger  will  fail  to 
arouse  contraction  of  a  muscle. 

Conductivity  in  Muscle. — In  an  ameba  or  other  simple  undifferentiated 
contractile  protoplasmic  unit  a  stimulus  applied  at  any  point  is  quickly 
transmitted  throughout  the  entire  mass.  Just  so  is  it  with  differentiated 
muscle.  A  stimulus  applied  at  any  point  of  a  muscle  will  quickly  be  propa- 
gated through  the  mass  as  far  as  there  is  protoplasmic  continuity.  In  cardiac 
muscle  and  in  smooth  muscle  there  is  uninterrupted  conduction  from  cell 
to  cell.  But  in  voluntary  muscle  each  fiber  is  physiologically  isolated  from 
its  neighbors.  When  a  voluntary  muscle  fiber  is  stimulated  either  at  the  ex- 
tremities or  at  its  middle,  the  effect  of  the  stimulus  quickly  passes  through 
the  entire  fiber,  whether  it  arouses  a  distinct  act  of  contraction  or  not. 

The  rate  at  which  conduction  takes  place  when  a  contraction  accom- 
panies it  has  been  carefully  measured  by  numerous  observers.  It  varies 
greatly  in  the  different  kinds  of  muscle,  from  two-tenths  of  a  meter  per  second 
in  the  rabbits'  ureter  (Engelmann)  to  ten  meters  per  second  in  the  voluntary 
muscles  of  man. 

SINGLE  MUSCLE  CONTRACTIONS. 

Characteristics  of  a  Single  Contraction. — The  Myogram. — The  con- 
traction of  a  muscle  in  response  to  a  single  effective  stimulus  of  short  dura- 
tion is  called  a  simple  muscle  contraction.  A  record  of  such  a  contraction 

33 


514  MUSCLE-NERVE    PHYSIOLOGY 

is  called  a  myogram.  The  character  of  the  myogram,  and  therefore  the  facts 
revealed  by  it,  are  dependent  on  whether  or  not  the  record  is  made  on  a  rapidly 
moving  recording  surface.  If  the  myogram  is  made  on  a  recording  surface 
that  is  standing  still,  then  it  shows  merely  the  extent  of  shortening  of  the 
muscle.  The  amount  of  shortening  for  a  given  muscle  will  depend  on  a  series 
of  conditions,  such  as  nutrition,  load,  temperature,  etc.,  all  of  which  will 
be  discussed  presently. 

When  the  record  is  made  on  a  rapidly  moving  drum  or  on  the  pendulum 
myograph,  it  is  revealed  that  the  simple  contraction  occupies  a  definite  per- 
iod of  time  with  well-marked  periods  or  phases.  Although  the  stimulus  may 
be  practically  instantaneous,  the  contraction  lasts  a  considerable  fraction  of  a 
second,  in  the  frog's  gastrocnemius  about  o.  i  of  a  second. 


FIG.  322  — Record  of  a  Simple  Contraction  of  the  Gastrocnemius  of  the  Frog.  Time 
in  o.oi  second.  Si,  Moment  of  stimulation.  Record  taken  on  a  rapid  drum  that  was 
provided  with  an  automatic  key. 

It  will  be  observed  that  after  the  stimulus  has  been  applied,  as  indicated 
by  the  vertical  line  St,  there  is  an  interval  before  contraction  commences. 
This  interval,  termed  the  latent  period,  when  measured  by  the  number  of  vi- 
brations of  the  tuning-fork  directly  beneath,  is  found  to  be  about  o.oi  of  a 
second.  The  latent  period  is  longer  in  some  muscles  than  in  others  and 
differs  also  according  to  the  condition  of  the  muscle  and  the  kind  of  stimulus 
employed.  During  the  latent  period  there  is  no  apparent  change  in  the 
muscle.  The  second  part  of  the  record  shows  the  contraction  phase  proper. 
The  lever  is  raised  by  the  sudden  shortening  of  the  muscle.  The  contrac- 
tion is  at  first  very  rapid,  but  then  progresses  more  slowly  to  its  maximum. 
It  occupies  on  an  average  o .  04  of  a  second  in  the  frog's  gastrocnemius.  The 
third  stage  is  the  relaxation  phase.  After  reaching  its  highest  point,  the  lever 
begins  to  descend,  in  consequence  of  the  elongation  of  the  muscle.  At- first 
the  fall  is  rapid,  but  it  then  becomes  more  gradual  until  the  lever  reaches  the 
abscissa  or  base  line,  when  the  muscle  has  attained  its  precontraction  length. 
The  stage  occupies  o .  05  of  a  second.  Usually  after  the  contraction  proper 
is  over  the  lever  oscillates  below  and  above  the  base  line  in  a  series  of  dimin- 
ishing waves,  the  elastic  rebound  following  movement  of  the  simple  contrac- 


CHANGE    IN    SHAPE    DURING    MUSCULAR    CONTRACTION  51$ 

tion.  These  are,  of  course,  wholly  passive  and  would  occur  equally  well  if 
we  should  lift  the  weight  to  the  height  of  the  contraction,  then  simply  let  it 
fall  while  taking  a  record. 

Change  in  Shape  during  Muscular  Contraction. — There  is  a  consider- 
able difference  of  opinion  as  to  the  mode  in  which  the  transversely  striated 
muscular  fibers  contract.  The  most  probable  account  is  that  the  contraction 
is  effected  by  an  approximation  of  the  constituent  parts  of  the  fibrils,  which, 
at  the  instant  of  contraction,  without  any  alteration  in  their  general  direction, 
become  closer,  flatter,  and  wider,  a  condition  which  is  rendered  evident  by 
the  approximation  of  the  transverse  striae  seen  on  the  surface  of  the  fasciculus, 
and  by  its  increased  breadth  and  thickness.  The  appearance  of  the  zigzag 
lines  into  which  it  was  supposed  the  fibers  are  thrown  in  contraction  is  due 
to  the  relaxation  of  a  fiber  which  has  been  recently  contracted  and  is  not 


FIG.  323.' — The  Microscopic  Appearances  During  a  Muscular  Contraction  in  the 
Individual  Fibrillae,  after  Engelmann.  i.  A  passive  muscle  fiber;  c  to  d  =  doubly  refractive 
discs,  with  median  disc  a  &  in  it;  h  and  g  are  lateral  discs;/ and  e  are  secondary  discs,  only 
slightly  doubly  refractive;  figure  on  right  same  fiber  in  polarized  light.  The  bright  part 
is  doubly  refracted,  black  ends  not  so.  2.  Transition  stage.  3.  Stage  of  entire  contrac- 
tion. In  each  case  the  right-hand  figure  represents  the  effect  of  polarized  light.  (Landois, 
after  Engelmann.) 

at  once  stretched  again  by  some  antagonist  fiber  or  whose  extremities  are 
kept  close  together  by  the  contractions  of  other  fibers.  The  contraction  is 
therefore  a  simple  and,  according  to  Edward  Weber,  a  uniform,  simultaneous, 
and  steady  shortening  of  each  fiber  and  its  contents.  What  each  fibril  or 
fiber  loses  in  length,  it  gains  in  thickness.  The  contraction  is  a  change  of 
form,  not  of  size;  it  is,  therefore,  not  attended  with  any  diminution  in  bulk 
from  condensation  of  the  tissue.  This  has  been  proved  for  entire  muscles 
by  making  a  mass  of  muscles,  or  many  fibers  together,  contract  in  a  vessel 
full  of  water,  with  which  -a  fine,  perpendicular,  graduated  tube  communi- 
cates. Any  diminution  of  the  bulk  of  the  contracting  muscle  would  be 
attended  by  a  fall  of  fluid  in  the  tube,  but  when  the  experiment  is  carefully 
performed,  the  level  of  the  water  in  the  tube  remains  the  same,  whether  the 
muscle  be  contracted  or  not. 

In  thus  shortening,  muscles  appear  to  swell  up,  becoming  rounder,  more 
prominent,  harder,  and  apparently  tougher.  But  this  hardness  of  muscle  in 
the  state  of  contraction  is  not  due  to  increased  firmness  or  condensation  of  the 
muscular  tissue,  but  to  the  increased  tension  to  which  the  fibers,  as  well  as 


MUSCLE-NERVE   PHYSIOLOGY 

their  tendons  and  other  tissues,  are  subjected  from  the  resistance  ordinarily 
opposed  to  their  contraction.  When  no  resistance  is  offered,  as  when  a 
muscle  is  cut  off  from  its  tendon,  not  only  is  no  hardness  perceived  during 
contraction,  but  the  muscular  tissue  is  even  softer  and  more  extensible  than 
in  its  ordinary  uncontracted  state.  During  contraction  in  each  fiber  it  is 
said  that  the  anisotropous  or  doubly  refractive  elements  become  less  refract- 
ive and  the  singly  refractive  more  so,  figure  321. 

Chemical  Changes  in  Contracting  Muscle. — i.  The  reaction  of  the 
muscle,  which  is  normally  alkaline  or  neutral,  becomes  decidedly  acid  during 
contraction,  from  the  development  of  sarcolactic  acid.  2.  The  muscle  gives 
out  carbon  dioxide  gas  and  takes  up  oxygen.  The  amount  of  the  carbon 
dioxide  given  out  does  not  appear  to  be  entirely  dependent  upon  the  oxygen 
taken  in,  and  so  doubtless  in  part  arises  from  some  other  source.  Muscle 
contracts  in  an  atmosphere  of  hydrogen,  showing  that  oxygen  is  present  in 
fixed  combination.  A  muscle,  however,  contracts  for  a  longer  time  in  an 
atmosphere  of  oxygen.  3.  Certain  imperfectly  understood  chemical  changes 
occur,  in  all  probability  connected  with  i  and  2,  in  which  glycogen  is  con- 
verted into  dextrose  and  the  latter  oxidized. 

Electrical  Changes  in  Contracting  Muscle. — Resting  muscles  un- 
injured in  the  body  have  a  uniform  potential,  i.e.,  are  isoelectric.  But  when 
removed  from  the  body  they  are  more  or  less  injured  and,  therefore,  show 
differences  of  electrical  potential  between  different  points  on  the  muscle, 
called  currents  of  injury  or  demarcation  currents.  It  is  necessary  to  use 
non-polarizable  and  not  metallic  electrodes  in  this  experiment,  as  other- 
wise there  is  no  certainty  that  the  whole  of  the  current  observed  is  com- 
municated from  the  muscle  itself  and  not  derived  from  the  metallic  elec- 
trodes and  arising  in  consequence  of  the  action  of  the  saline  juices  of  the 
tissues  upon  them.  Non-polarizable  electrodes  are  usually  some  modifica- 
tion of  Du  Bois  Reymond's  apparatus,  which  consists  of  a  cylinder  filled  with 
china  clay  moistened  with  saline  solution,  part  of  which  projects  as  a 
drawn-out  point  for  contact  with  the  muscle.  The  rest  of  the  cylinder  is 
filled  with  a  saturated  solution  of  zinc  sulphate  into  which  dips  a  well-amal- 
gamated piece  of  zinc  connected  by  wire  with  the  galvanometer.  In  this  way 
the  zinc  sulphate  and  the  sodium  chloride  form  a  non-polarizable  conductor 
between  the  zinc  and  the  live  muscle.  Recently  Porter  has  devised  a  boot- 
shaped  clay  electrode  that  is  burned,  and  hence  has  the  immense  advan- 
tage of  permanency. 

Currents  of  Injury,  or  Demarcation  Currents. — If  a  segment  is  cut 
out  of  a  living  gastrocnemius,  its  cut  ends  present  regions  of  maximal  injury. 
Such  a  preparation  is  called  a  muscle  prism. 

If  the  points  on  the  surface  of  a  muscle  prism  be  connected  with  the  gal- 
vanometer by  non-polorizable  electrodes,  it  will  be  found  that  the  currents 
pass  from  point  to  point,  as  shown  in  the  diagram,  figure  324. 


HEAT    PRODUCED    IN    A    SIMPLE    CONTRACTION 


517 


The  strongest  currents  pass  from  the  equator  to  a  point  representing  the 
middle  of  the  cut  ends;  currents  also  pass  from  points  nearer  the  equator  to 
those  more  remote,  but  not  from  points  equally  distant,  which  are  isoelectric 
points.  The  cut  ends  are  always  negative  to  the  equator.  The  currents  are 
in  all  probability  due  to  chemical  changes  going  on  in  the  muscles  at  the  in- 
jured ends. 

Action  Currents. — When  a  muscle  is  made  to  contract  the  demar- 
cation current  undergoes  a  sharp  decrease,  as  shown  by  the  deflection  of  the 
galvanometer  needle,  which  swings  back  in  the  direction  of  equilibrium. 
This  deflection,  originally  called  the  current  of  negative  variation,  has  been 
shown  to  be  due  to  the  processes  going  on  in  the  muscle  during  contraction, 
and  is  therefore  more  properly  called  the  action  current.  It  occurs  where  no 
previous  demarcation  current  exists. 

For  the  study  of  the  action  current  the  capillary  electrometer  is  very  con- 
venient. The  hearts  of  cold-blooded  animals,  because  of  their  slow  con- 
traction, serve  well  for  demonstration.  The  muscle  contraction  passes  over 


FIG.  324. — Diagram  of  the  Currents  in  a  Muscle  Prism.     (Du  Bois  Reymond.) 

the  ventricle  in  the  form  of  a  wave,  the  electric  potential  of  the  muscle  chang- 
ing as  it  becomes  active  or  passive.  For  any  two  points  on  the  heart  muscle, 
therefore,  there  will  be  two  changes  of  potential,  the  active  part  first  becom- 
ing negative  to  the  inactive,  and  then,  as  the  wave  passes  down  and  the  in- 
active part  becomes  active,  the  current  is  reversed.  This  is  known  as  a 
diphasic  current. 

In  certain  fishes  definite  electrical  organs  exist,  organs  which  are  derived 
from  muscle-like  tissues  and  which  may  be  regarded  morphologically  as  mus- 
cles highly  specialized  for  the  production  of  energy  in  the  form  of  electricity. 

Heat  Produced  in  a  Simple  Contraction. — Becquerel  and  Breschet 
found,  with  the  thermo-multiplier,  about  0.5°  C.  of  heat  produced  by  each 
forcible  contraction  of  a  man's  biceps;  and  when  the  actions  were  long  con- 
tinued, the  temperature  of  the  muscle  increased  i°  C.  In  the  frog's  muscle 
a  considerable  number  of  contractions  have  been  found  to  produce  an  ele- 
vation of  temperature  equal  on  an  average  to  less  than  o.  2°  C.,  while  a  single 
contraction  produces,  according  to  R.  Heidenhain,  from  0.001°  to  0.005°  C. 


Si* 


MUSCLE-NERVE   PHYSIOLOGY 


One  gram  of  frog's  muscle  will  produce  in  a  single  maximal  contraction  about 
0.003  calorie  or  the  equivalent  of  128  gramcentimeters  of  work  energy  (since 
i  calorie  =0.4267  kilogrammeter  of  work).  The  cause  of  the  rise  of  tempera- 
ture is  the  increased  chemical  activity  at  the  time  of  contraction.  As  we 
have  already  seen,  in  the  chapter  on  Animal  Heat,  muscles  have  the  power  of 
producing  heat  even  when  not  contracting,  i.e.,  changing  shape. 

The  amount  of  heat  energy  developed  during  a  single  contraction  will 
vary  sharply  according  to  the  tension  under  which  the  muscle  contracts. 
The  heat  production  follows  closely  the  energy  of  work  produced,  and 
apparently  obeys  the  same  laws. 


FIG.  325. — Figure  for  Work  Energy,  Showing  Height  of  the  Contraction  of  the  Gastroc- 
nemius  of  the  Frog  with  Loads  Increased  by  Ten  Grams  at  a  Time. 

The  Work  Energy  Liberated  by  a  Simple  Muscle  Contraction.— 

When  a  muscle  contracts  against  a  resistance  and  a  load  is  moved,  work 
energy  is  liberated.  In  fact,  the  liberation  of  work  energy  and  heat  energy 
are  the  specific  functions  of  the  muscles  among  the  warm-blooded  animals. 
A  frog's  gastrocnemius  weighing  i  gram  and  loaded  with  50  grams  will 
contract  from  0.5  to  0.6  cm.;  i.e.,  will  do  25  to  30  gramcentimeters  of  work 

TABLE  SHOWING  THE  RELATION  BETWEEN  LOAD  AND  WORK. 


1 

Load  or  tension. 

Height  lifted.                         Work  done. 

Grams. 

Centimeters. 

Gramcentimeters. 

0 

I  .2 

0 

40 

0.8 

32 

80 

o-5 

40 

120 

0.4 

48 

160 

0.2 

32 

2OO 

O.  I 

20 

240 

o  .0 

0 

EFFECT    OF    THE    STRENGTH    OF    STIMULUS  519 

for  each  simple  contraction.  The  amount  of  work  done  is  intimately  associ- 
ated with  the  tension  under  which  the  muscle  contracts.  As  the  tension  in- 
creases from  no  load  up  to  100  or  150  grams  (for  a  i-gram  muscle),  the  work 
increases.  But  as  the  tension  continues  to  increase,  the  work  falls  off  until 
a  point  is  reached  at  which  the  load  is  not  lifted  at  all. 

CONDITIONS  WHICH  AFFECT  THE   IRRITABILITY  OF 

THE  MUSCLE  AND  THE  CHARACTER  OF  THE 

CONTRACTION. 

There  are  a  number  of  conditions  which  influence  both  the  irritability 
of  a  muscle  and  the  power  and  character  of  its  contractions.  Irritability 
and  contractility  may  vary  independently,  but  as  a  rule  any  condition  which 
decreases  one  also  decreases  the  other.  The  most  important  of  these  con- 
ditions are:  relation  of  the  muscle  to  the  central  nervous  system,  condition 
of  nutrition,  stimulus,  temperature,  fatigue,  drugs,  disease,  etc. 

Effect  of  the  Strength  of  Stimulus. — A  strength  of  current  that  is  just 
sufficient  to  give  the  contraction  of  a  muscle  is  called  a  minimal  stimulus. 


FIG.  326. — Contraction  of  the  Gastrocnemius  Under  the  Influence  of  Variation  of 
Strength  of  Stimulus.  The  muscle  was  stimulated  by  Petzold's  inductorium,  graduated  to 
show  units  of  current.  The  figures  6,  7,  8,  9,  10,  etc.,  indicate  relative  strength  of  stimulus. 

This  is  a  comparatively  weak  induction  current,  one  which  can  scarcely  be 
detected  by  the  tip  of  the  tongue.  As  the  strength  of  the  current  is  very 
gradually  increased,  the  height  of  the  contraction  curve  increases  until  the 
maximal  stimulus  is  reached,  which  produces  a  contraction  of  an  amplitude 
beyond  which  no  increase  occurs  even  though  the  strength  of  the  stimulus  be 
multiplied  many  fold.  The  range  between  the  strengths  of  the  minimal  and 
maximal  stimuli  is  very  restricted  indeed.  The  absolute  strength  of  a  mini- 


520  MUSCLE-NERVE    PHYSIOLOGY 

mal  stimulus  varies  exceedingly  for  a  given  muscle,  depending  on  its  degree 
of  irritability.  This  narrow  range  between  minimal  and  maximal  stimuli 
serves  as  a  convenient  means  for  detecting  the  variations  in  irritability.  One 
should  count  on  a  continued  decrease  in  irritability  in  isolated  muscles,  hence 
should  choose  a  supramaximal  stimulus  for  all  such  preparations  when  other 
conditions  surrounding  the  muscle  are  under  investigation. 

The  Influence  of  Repeated  Activity. — The  irritability  of  muscle  is 
decreased  by  undue  functional  activity.  The  cause  of  the  diminished  ir- 
ritability is  twofold:  when  a  muscle  contracts,  part  of  its  substance  is  ex- 
pended, part  of  its  store  of  nutriment  is  exhausted,  and  it  cannot  contract  so 


FIG.  327. — Contractions  of  the  Gastrocnemius  Muscle  to  Show  Fatigue.     The  numbers 
printed  on  the  figure  indicate  the  contractions  in  the  series  which  is  recorded.     (Lee.) 

vigorously  again  until  the  loss  is  made  up.  To  this  extent  fatigue  has  much 
the  same  effect  as  cutting  off  or  diminishing  the  blood  supply.  The  other 
cause  for  the  diminution  of  irritability  is  the  accumulation  of  poisonous  prod- 
ucts in  the  muscle,  substances  generated  during  contraction,  probably  sar- 
colactic  acid  chiefly.  In  a  living  animal  these  poisonous  products  exert  their 
influence  not  only  upon  the  muscle  or  muscles  immediately  concerned  in 
contraction,  but  upon  the  musculature  of  the  body  generally,  and  the  effect 
remains  until  they  are  eliminated  from  the  body.  Massage  of  the  muscles 
increases  the  passage  of  waste  products  into  the  general  blood  stream  and  the 
rapidity  of  their  elimination. 

In  the  first  few  simple  contractions,  repeated  in  series,  there  is  an  increase 
in  the  amplitude  of  the  contractions  resulting  in  the  phenomenon  known 
as  staircase  contractions  or  ' '  Treppe. "  This  stage  is  followed  by  a  period 
of  sustained  contractions,  and  this  finally  by  a  diminishing  series  of  ampli- 
tudes until  the  muscle  fails  to  respond.  After  a  few  minutes'  rest  a  muscle 
will  again  give  strong  contractions,  but  only  for  a  brief  series. 

If  the  time  of  the  simple  contractions  is  measured,  it  will  be  found,  figure 
327,  that  not  only  is  the  amplitude  decreased,  but  the  duration  is  greatly 


THE  INFLUENCE  OF  TEMPERATURE  521 

increased  as  the  contractions  are  repeated.  The  latent  period  changes  very 
little.  The  contraction  phase  is  considerably  prolonged,  but  the  relaxation 
phase  is  very  greatly  increased.  As  fatigue  progresses,  the  total  time  of  the 
simple  contraction  may  be  two  or  three  times  longer  than  the  normal.  The 
ability  of  the  muscle  to  do  work  falls  off  rapidly,  of  course;  and  the  greater 
the  load  during  the  time  fatigue  is  coming  on,  the  more  quickly  complete 
fatigue  approaches. 

The  Influence  of  Temperature. — The  irritability  of  muscle  is  in- 
creased by  raising  its  temperature  slightly  above  that  of  the  animal  from 
which  it  has  been  taken,  while  it  is  decreased  by  cooling.  If,  however,  the 
temperature  be  raised  too  high  (40°  C.  for  frog,  50°  C.  for  mammal),  the 
muscle  enters  into  a  condition  of  heat  rigor  and  its  irritability  is  forever  lost. 


FIG.  328. — Contractions  of  ihe  Gastrocnemius  Muscle  to  Show  the  Influence  of 
Temperature  on  the  Amplitude  of  the  Contractions.  At  40°  C.  the  muscle  has  begun  to 
pass  into  rigor  mortis,  yet  is  able  to  give  short  contractions.  The  steps  on  the  curve  of  rigor 
at  the  right  occur  at  temperatures  of  41°,  42°,  and  43°  C. 

After  cooling,  unless  the  cold  be  too  severe  and  prolonged,  the  irritability  re- 
turns as  the  temperature  is  raised.  A  series  of  vertical  records  of  simple  con- 
tractions beginning  at  room  temperature  and  decreasing  with  a  contraction 
at  each  fall  of  one  degree  reveals  the  fact  that  the  amplitude  falls  off  slowly 
until  a  temperature  of  12°  to  10°  C.  is  reached,  then  as  gradually  increases 
until  4°  to  2°  C.,  after  which  the  amplitude  drops  off  sharply  to  about  -  i°  C. 
However,  this  phenomenon  is  partly  one  of  irritability,  since  a  very  strong 
stimulus  will  produce  a  vigorous  contraction  until  the  muscle  begins  to 
freeze.  If  at  the  freezing  temperature  the  muscle  be  slowly  and  carefully 
increased  in  temperature  it  will  recover  from  the  effects  of  the  cooling  with- 
out apparent  injury,  and  will  give  a  reverse  series  to  the  one  obtained  by 
decreasing  the  temperature.  As  the  increase  of  temperature  is  continued 
above  room  temperature  the  amplitude  of  the  contractions  very  greatly 
increases  (also  the  elasticity),  reaching  a  maximum  in  the  frog's  gastroc- 
nemius  at  about  35°  to  36°  C.  The  amplitude  sharply  decreases  above 


522  MUSCLE-NERVE    PHYSIOLOGY 

35°  C.  up  to  37°  to  38°  C.,  where  heat  rigor  begins  and  the  muscle  per- 
manently shortens.  Heat  rigor  is  usually  complete  at  40°  to  41°  C.  A 
muscle  cannot  recover  its  irritability  after  heat  rigor  has  set  in  strongly. 

If  the  time  of  the  contraction  is  measured  at  different  temperatures  it  will 
be  found  to  be  greatly  delayed  at  2°  to  4°  C.,  and  very  much  quicker  than  nor- 
mal at  33°  to  35°  C.  As  in  fatigue,  the  effect  falls  chiefly  on  the  contraction 
and  relaxation  phases  and  only  slightly  on  the  latent  period.  The  latent 
period  is  more  sharply  influenced  by  temperature  than  by  fatigue. 


FIG.  329. — Influence  of  Temperature  on  the  Duration  of  the  Contraction  of  the  Frog's 

Gastrocnemius. 

Influence  of  Blood  Supply. — In  the  normal  human  muscle  there  is  a 
delicately  balanced  vaso-motor  mechanism  by  which  the  amount  of  blood 
flowing  through  a  muscle  is  immediately  increased  when  the  muscle  is  in  con- 
traction. This  blood  stream  is  of  course  carrying  nutritive  materials  to  the 
muscle  and  taking  away  wastes.  If  the  blood  supply  to  a  muscle  is  cut  off, 
then  the  muscle  can  only  draw  on  its  stored  supply  of  potential  energy,  which 
in  active  contraction  is  sooner  or  later  exhausted.  Under  such  conditions 
the  muscle  increases  in  irritability  for  a  few  minutes  and  then  gradually  loses 
both  its  irritability  and  its  power  to  contract.  Even  mammalian  muscles 
have  been  kept  alive  and  normal  in  their  activity  for  several  hours  by  irri- 


EFFECT  OF  RATE  OF  STIMULATION  523 

gating  them  with  defibrinated  and  aerated  blood  (von  Frey).  Mammalian 
muscles  will  remain  irritable  for  30  minutes,  or  longer  if  cooled,  after  being 
shut  off  from  their  blood  supply  and  isolated  from  the  body,  but  both  irrita- 
bility and  contractility  soon  disappear  entirely. 

Effect  of  Nerve  Supply. — The  voluntary  or  skeletal  muscle  normally 
contracts  in  the  body  only  when  stimulated  through  its  motor  nerve.  If  the 
motor  nerve  is  severed,  the  muscle  is  cut  off  from  its  normal  source  of  activity, 
hence  will  undergo  the  changes  resulting  from  disuse,  which  will  be  presently 
discussed.  Aside  from  this,  it  is  held  by  most  observers  that  there  are  dis- 
tinct nutritive  or  trophic  nerves  which  exercise  a  controlling  influence  over 
the  growth,  development,  and  general  nutritive  processes  going  on  in  muscle. 

When  a  motor  nerve  is  cut,  the  muscle  at  first  exhibits  heightened  irrita- 
bility to  all  forms  of  stimuli.  In  a  couple  of  weeks  it  decreases  in  its  power 
to  respond  to  rapidly  changing  stimuli  like  induced  currents.  It  responds 
more  readily  to  mechanical  shocks  and  to  galvanic  currents  for  six  or  seven 
weeks,  then  gradually  loses  the  power  of  contracting  through  as  many  months. 
The  changes  are  due  to  protoplasmic  degeneration.  It  is  not  clear  in  what 
degree  these  changes  are  due  to  loss  of  trophic  nerve  influence,  to  inactivity, 
and  to  changes  in  nutritive  conditions.  Since  degeneration  occurs  when 
the  vascular  supply  is  maintained,  it  would  seem  that  the  nutritive  condition 
must  be  chargeable  to  one  or  the  other  of  the  first  two  factors,  probably  to 
both. 

Use  of  muscle  increases  its  power  and  also  its  irritability.  A  properly 
regulated  exercise  is  well  known  to  contribute  to  the  health  and  development 
of  muscles.  In  cases  of  paralysis,  mechanical  or  electrical  stimulation  is 
applied  directly  to  the  muscle  in  an  effort  to  supply  artificial  exercise  until  the 
nerves  are  regenerated  and  motor  connections  re-established.  If  such  stim- 
ulation is  not  applied,  the  muscle  degenerates  from  disuse  and  loses  its 
irritability  often  before  the  nerves  regenerate. 

The  Effect  of  Drugs. — Drugs  affect  the  irritability  of  muscle,  some 
augmenting,  others  depressing  it.  Voluntary  muscle,  which  does  not  ordina- 
rily contract  except  when  stimulated,  can  be  made  so  irritable  by  certain 
salts  that  it  contracts  automatically  like  heart  muscle,  and  the  converse. 
Ether,  chloroform,  etc.,  anesthetize  muscle  just  as  they  do  nerve,  suppressing 
both  irritability  and  contractility.  Suprarenal  extract  increases  the  ampli- 
tude of  contraction,  as  do  also  caffeine,  digitalis,  nicotine,  and  others.  Ver- 
atrine  is  well  known  greatly  to  prolong  the  relaxation  phase  of  the  simple 
contraction  without  materially  affecting  the  contraction  phase  or  the  latent 
period. 

TETANIC  AND  VOLUNTARY  MUSCULAR  CONTRACTIONS. 

Effect  of  Rate  of  Stimulation. — If  we  stimulate  the  muscle:nerve  prep- 
aration with  two  induction  shocks,  one  immediately  after  the  other,  when  the 


524  MUSCLE-NERVE    PHYSIOLOGY 

point  of  stimulation  of  the  second  one  corresponds  to  the  crest  of  the  con- 
traction of  the  first,  a  second  curve,  figure  330,  will  occur,  which  will  commence 
near  the  highest  point  of  the  first  and  will  rise  nearly  as  much  higher,  so  that 
the  sum  of  the  height  of  the  two  curves  almost  exactly  equals  twice  the  height 
of  the  first.  This  phenomenon  is  called  summation.  If  a  third  and  fourth 
shock  be  passed,  a  similar  effect  will  ensue,  and  curves  one  above  the  other 
will  be  traced,  the  third  being  slightly  lower  than  the  second,  and  the  fourth 
than  the  third.  If  a  continuous  series  of  shocks  occur,  however,  the  lever 
after  a  time  ceases  to  rise  any  further,  and  the  contraction,  which  has  reached 


FIG.  330. — Tracing  of  a  Double  Muscle  Curve.  To  be  read  from  left  to  right.  While 
the  muscle  was  engaged  in  the  first  contraction  (whose  complete  course,  had  nothing 
intervened,  is  indicated  by  the  dotted  line),  a  second  induction  shock  was  thrown  in,  at 
such  a  time  that  the  second  contraction  began  just  as  the  first  was  beginning  to  decline. 
The  second  curve  is  seen  to  start  from  the  first,  as  does  the  first  from  the  base  line.  (M. 
Foster.) 

its  maximum,  is  maintained.  The  condition  which  ensues  is  called  Tetanus. 
A  tetanus  is  really  a  summation  of  contractions,  but  unless  the  stimuli  become 
very  rapid  indeed,  the  muscle  will  still  be  in  a  condition  of  vibratory  contrac- 
tion and  not  of  unvarying  contraction. 

If  the  shocks,  however,  be  repeated  at  very  short  intervals,  varying,  in  the 
frog,  from  eighteen  to  thirty  per  second,  the  muscle  contracts  to  its  utmost 
at  once  and  continues  at  its  maximum  contraction  for  some  time.  The 
lever  rises  almost  perpendicularly  and  then  describes  a  straight  line,  figure 
331,  c.  The  rate  of  stimulation  required  increases  with  the  rapidity  of  the 
simple  contraction.  If  the  stimuli  are  not  so  rapid  the  line  of  maximum  con- 
traction becomes  wavy,  indicating  a  tendency  of  the  muscle  to  relax  during 
the  intervals  between  the  stimuli,  figure  331,  b.  As  the  muscle  becomes 
fatigued,  a  less  rapid  rate  of  stimulation  is  required  to  produce  a  complete 
tetanus,  owing  to  the  prolongation  of  the  relaxation  period  in  such  a  muscle. 
The  height  of  the  contraction,  however,  is  lessened.  This  condition  of  pro- 
longed relaxation  is  known  as  contracture. 

Co-ordinated  Muscular  Contractions. — In  the  human  body  the  skel- 
etal muscles  contract  only  on  stimulation  through  their  motor  nerves;  i.e., 
under  the  influence  of  nerve  impulses  that  have  their  origin  in  the  central 


CO-ORDINATED    MUSCULAR    CONTRACTIONS 


525 


nervous  system.  Such  motor  impulses  may  arise  through  reflexes,  through 
automatic  activity  of  the  nerve  center,  or  by  higher  cerebral  origin  associated 
with  conscious  psychic  effort.  In  either  case  the  apparatus  consists  of  one 
or  more  central  neurones,  an  anterior-horn  motor  cell,  and  the  muscle  itself. 
Conscious  or  voluntary  effort  may  be  taken  as  a  type. 

Simple  contractions  are  possible  to  human  muscles,  but  undoubtedly 
tetanic  contractions  are  the  rule.     If  one  holds  the  arm  out  at  right  angles  to 


FIG.  331. — a,  Frog's  gastrocnemius  muscle  stimulated  with  four  induction  shocks  per 
second,  showing  complete  relaxation  between  stimuli;  b,  same  muscle  stimulated  eight  times 
per  second,  showing  partial  relaxation  between  stimuli  (incomplete  tetanus);  c,  same 
muscle  stimulated  twelve  times  per  second,  showing  development  of  an  almost  complete- 
tetanus. 

the  trunk,  the  movement  requires  the  continuous  or  tetanic  contraction  of  the 
deltoid  and  the  series  of  extensor  muscles.  If  the  arm  is  retained  in  the 
extended  position  long  enough,  extreme  fatigue  is  felt  and  presently  one  can 
no  longer  maintain  the  position.  Yet,  if  the  muscles  involved  are  immediately 
stimulated  directly  with  an  electric  current,  they  contract,  showing  that  such 
exhaustion  as  exists  is  not  wholly  due  to  the  muscle. 

Mosso's  ergograph  was  devised  for  the  specific  purpose  of  studying  the 
character  of  fatigue  of  voluntary  effort.  This  apparatus  is  adapted  to  the 


526  MUSCLE-NERVE   PHYSIOLOGY 

study  of  the  fatigue  of  the  flexors  of  the  middle  finger  or,  in  the  newer  in- 
strument devised  by  Storey,  to  the  abductor  of  the  index  finger.  Numerous 
studies  have  shown,  apparently,  that  the  fatigue  of  voluntary  effort  involves, 
first,  the  nervous  apparatus  and,  later,  the  muscle;  that  the  muscle  still 
retains  a  considerable  reserve  of  energy  when  the  apparatus  as  a  whole  is 
exhausted. 

Muscle  in  Rigor  Mortis. — After  the  muscles  of  the  dying  body  have 
lost  their  irritability  or  capability  qf  being  excited  to  contraction  by  the  ap- 
plication of  a  stimulus,  they  spontaneously  pass  into  a  state  of  shortening 
apparently  identical  in  effect  with  that  which  ensues  during  life.  It  affects 
all  the  muscles  of  the  body,  and,  when  external  circumstances  do  not 
prevent  it,  commonly  fixes  the  limbs  in  that  which  is  their  natural  posture  of 
equilibrium  or  rest.  From  the  simultaneous  contraction  of  all  the  muscles 
of  the  trunk,  a  general  stiffening  of  the  body  is  produced,  which  constitutes 
the  rigor  mortis  or  postmortem  rigidity. 

When  this  condition  has  set  in,  the  muscle  becomes  acid  in  reaction  (due  to 
development  of  sarcolactic  acid),  gives  off  carbonic  acid  in  great  excess, 
diminishes  in  volume  slightly,  becomes  shortened  and  opaque,  its  substance  sets 
in  a  firm  coagulation.  Rigor  mortis  comes  on  much  more  rapidly  in  death 
after  muscular  activity,  and  is  hastened  by  warmth. 

The  immediate  cause  of  rigor  mortis  seems  to  be  a  chemical  one,  namely, 
the  coagulation  of  the  muscle  plasma.  We  may  distinguish  three  main 
stages:  i,  gradual  coagulation;  2,  contraction  of  the  coagulated  muscle 
clot  (myosin)  and,  3,  squeezing  out  of  muscle  serum.  During  the  first  stage, 
restoration  is  possible,  by  the  circulation  of  arterial  blood  through  the  muscles, 
and  even  when  the  second  stage  has  set  in,  vitality  may  be  restored  by  dis- 
solving the  coagulum  of  the  muscle  by  salt  solution  and  by  passing  arterial 
blood  through  the  vessels.  After  the  second  stage  is  advanced,  recovery  is 
impossible. 

It  has  been  noticed  that  the  relaxation  in  muscles  after  rigor  sometimes 
occurs  too  quickly  to  be  caused  by  putrefaction.  The  suggestion  that  in 
such  cases  the  relaxation  is  due  to  a  ferment  action  is  very  plausible. 
Subjecting. fresh  muscle  to  the  action  of  heat  (50°  to  60°  C.)  or  immersing 
it  in  distilled  water  causes  a  similar  coagulation  to  that  of  rigor  mortis. 
The  former  is  known  as  heat  rigor,  and  the  latter  as  water  rigor. 

The  muscles  are  not  affected  simultaneously  by  rigor  mortis.  It  affects 
the  neck  and  lower  jaw  first;  next,  the  upper  extremities,  extending  from 
above  downward;  and,  lastly,  reaches  the  lower  limbs.  In  some  rare  in- 
stances only  it  affects  the  lower  extremities  before  or  simultaneously  with 
the  upper  extremities.  It  usually  ceases  in  the  order  in  which  it  begins: 
first  at  the  head,  then  in  the  upper  extremities,  and  lastly  in  the  lower  ex- 
tremities. It  never  ordinarily  commences  earlier  than  ten  minutes,  and 
never  later  than  seven  hours  after  death;  and  its  duration  is  greater  in  pro- 


MUSCLE    IN    RIGOR    MORTIS  527 

portion  to  the  lateness  of  its  accession.     Heat  is  developed  during  the  passage 
of  a  muscular  fiber  into  the  condition  of  rigor  mortis. 

Since  rigidity  does  not  ensue  until  muscles  have  lost  the  capacity  of  being 
excited  by  external  stimuli,  it  follows  that  all  circumstances  which  cause  a 
speedy  exhaustion  of  muscular  irritability  induce  an  early  occurrence  of  the 
rigidity,  while  conditions  by  which  the  disappearance  of  the  irritability  is 
delayed  are  succeeded  by  a  tardy  onset  of  the  rigidity  of  rigor.  This  is  the 
explanation  of  its  speedy  occurrence,  and  equally  speedy  departure,  in  the 
bodies  of  persons  exhausted  by  chronic  diseases;  and  its  tardy  onset  and  long 
continuance  after  sudden  death  from  acute  diseases.  In  some  cases  of 
sudden  death  from  lightning,  violent  injuries,  or  paroxysms  of  passion,  rigor 
mortis  has  been  said  not  to  occur  at  all;  but  this  is  not  always  the  case.  It 


FIG.  332. — Curve  of  Shortening  of  the  Gastrocnemius  Muscle  of  the  Frog,  During  Heat 
Rigor.     The  numbers  indicate  degrees  centigrade. 

may,  indeed,  be  doubted  whether  there  is  really  a  complete  absence  of  the 
postmortem  rigidity  in  any  such  cases;  for  the  experiments  of  Brown- 
Sequard  make  it  probable  that  the  rigidity  may  supervene  immediately 
after  death,  and  then  pass  away  with  such  rapidity  as  to  be  scarcely 
observable. 

The  occurrence  of  rigor  mortis  is  not  prevented  by  the  previous  existence 
of  paralysis  in  a  part,  provided  the  paralysis  has  not  been  attended  with  very 
imperfect  nutrition  of  the  muscular  tissue. 

The  rigidity  affects  the  involuntary  as  well  as  the  voluntary  muscles, 
whether  they  be  constructed  of  striped  or  unstriped  fibers.  The  rigidity 
of  involuntary  muscles  with  striped  fibers  is  shown  in  the  contraction  of 
the  heart  after  death.  The  contraction  of  the  muscles  with  unstriped  fibers 
is  shown  by  an  experiment  of  Valentin,  who  found  that  if  a  graduated  tube 
be  connected  with  a  portion  of  intestine  taken  from  a  recently  killed  animal, 


MUSCLE-NERVE   PHYSIOLOGY 


and  the  intestine  be  tied  at  the  opposite  end,  and  filled  with  water,  the  water 
will  in  a  few  hours  rise  to  a  considerable  height  in  the  tube,  owing  to  the  con- 
traction of  the  intestinal  walls.  It  is  still  better  shown  in  the  arteries,  of 
which  all  that  have  muscular  coats  contract  after  death,  and  thus  present  the 
roundness  and  cord-like  feel  of  the  arteries  of  a  limb  lately  removed  or  those 
of  a  body  recently  dead.  Subsequently  they  relax,  as  do  all  the  other  mus- 
cles, and  feel  lax  and  flabby  and  lie  as  if  flattened  and  with  their  walls 
nearly  in  contact. 

Muscular  Metabolism  During  Contraction. — The  question  of  the 
metabolism  of  muscle  both  in  a  resting  and  in  an  active  condition  has  for 
many  years  occupied  the  attention  of  physiologists.  It  cannot  be  said  even 
now  to  be  thoroughly  understood.  Most  of  the  facts  with  reference  to  the 
subject  have  been  already  mentioned.  We  may  shortly  recapitulate  them 
here:  First,  muscle  during  rest  absorbs  oxygen  and  gives  out  carbon 
dioxide.  This  has  been  shown  by  an  analysis  of  the  gases  of  the  blood  going 
to  and  leaving  muscles.  During  activity,  e.g.,  during  tetanus,  the  same  inter- 
change of  gases  takes  place,  but  the  quantities  of  the  oxygen  absorbed  and 
of  the  carbon  dioxide  given  up  are  increased,  and  the  proportion  between 
them  is  altered  thus: 


Venous  blood. 

O2,  less  than  in  arterial 
blood. 

Co2,  more  than  in  arterial 
blood 

Of  restin°r  muscle 

9  per  cent 

6.71  per  cent 

Of  active  muscle               .  . 

12   26  per  cent 

10.79  per  cent. 

There  is,  then,  a  greater  proportion  of  carbon  dioxide  produced  in  muscle 
during  activity  than  during  rest. 

During  rigor  mortis  there  is  also  an  increased  production  of  carbon 
dioxide. 

Second,  muscle  during  rest  produces  nitrogenous  crystallizable  sub- 
stances, such  as  creatin,  from  the  metabolism  which  is  constantly  going  on 
in  it  during  life;  in  addition  there  are  formed,  in  all  probability,  sarcolactic 
acid  and  other  non-nitrogenous  matters. 

During  activity  the  nitrogenous  substances,  such  as  creatin,  undergo 
very  slight,  if  any,  increase  above  the  amount  produced  during  rest — but 
the  sarcolactic  acid  is  distinctly  increased.  The  glycogen  stored  in  the 
muscle  is  gradually  converted  into  dextrose  and  the  latter  oxidized  to 
furnish  the  energy  developed  in  the  contraction. 


CONTRACTION    IN    INVOLUNTARY    MUSCLE    AND    IN    CILIA          529 

During  rigor  mortis  the  sarcolactic  acid  is  increased,  and  in  addition 
myosin  is  formed. 

From  these  data  it  is  assumed  that  the  processes  which  take  place  in 
resting  and  active  muscles  are  somewhat  different,  at  any  rate  in  degree, 
from  actively  contracting  muscle.  Also,  there  are  obtained  an  increased 
amount  of  heat  and  mechanical  work;  potential  energy  is  converted  into 
kinetic  energy. 

THE  TYPE  OF  CONTRACTION  IN  INVOLUNTARY 
MUSCLE  AND  IN  CILIA. 

Cardiac  Muscle. — Some  detail  concerning  the  action  of  cardiac  muscle 
has  already  been  given  in  connection  with  the  chapter  on  Circulation. 
As  compared  with  the  activity  of  skeletal  muscle,  cardiac  muscle  differs  most 
strikingly  in  that  it  is  automatic.  A  strip  of  heart  muscle  taken  from  any 
part  of  the  heart,  under  proper  conditions,  gives  off  a  series  of  contrac- 
tions, whether  it  receives  any  special  stimulus  or  not,  whereas  we  have  just 
found  that  skeletal  muscle  under  similar  conditions  remains  quiet  unless 
stimulated  in  some  special  way.  The  fibers  of  skeletal  muscle  are  more  or 
less  physiologically  isolated  from  each  other,  and  one  fiber  may  contract  with- 
out involving  contractions  of  the  others.  Cardiac  muscle,  on  the  other  hand, 
when  stimulated  at  any  point  conducts  the  change  produced  throughout  the 
continuity  of  the  mass.  Cardiac  muscle  contractions  are  influenced -by 
tension,  temperature,  fatigue,  etc.,  apparently,  in  the  same  way  as  skeletal 
muscle. 

When  the  contraction  occurs  it  is  always  maximal.  The  actual  am- 
plitude of  the  contraction  is  dependent  on  the  condition  of  nutrition  of  the 
cardiac  muscle.  If  the  contractions  are  at  a  rapid  rate  they  will  be  relatively 
of  less  amplitude.  If  an  extra  contraction  is  induced  in  an  automatic  series, 
so  that  the  interval  between  two  contractions  is  similar,  then  the  amplitude 
will  be  correspondingly  reduced.  Such  an  extra  contraction  is  followed  by 
a  delayed  automatic  contraction,  the  phenomenon  of  compensatory  pause. 
The  contractions  in  cardiac  muscle  are  simple  contractions.  In  fact,  it  is  said 
to  be  impossible  to  produce  a  tetanus  except  in  certain  invertebrate  hearts. 
This  possibility  depends  upon  the  fact  that  during  the  time  of  a  single  con- 
traction there  is  a  certain  interval  between  the  beginning  and  the  crest  of  the 
contraction,  figure  174,  in  which  the  heart  muscle  is  not  irritable.  This  is 
known  as  the  refractory  phase. 

The  duration  of  the  contraction  of  heart  muscle  is  much  greater  than 
the  contraction  of  skeletal  muscle.  The  total  time  of  a  contraction  in  a  frog's 
gastrocnemius  is  o .  i  of  a  second,  while  the  time  of  a  contraction  of  the  ven- 
tricle in  the  same  animal  is  at  least  o .  7  to  o .  8  of  a  second.  In  the  terrapin's 
cardiac  muscle  the  time  of  a  contraction  is  over  a  second,  but  in  the  warm- 

34 


530 


MUSCLE-NERVE   PHYSIOLOGY 


blooded  cardiac  muscle  the  time  is  shorter,  perhaps  from  0.4  to  0.5  of  a 
second  for  the  human  ventricular  muscle. 

Smooth  Muscle. — The  physiology  of  smooth  muscle  has  been  given 
to  some  extent  in  previous  chapters,  particularly  in  connection  with  the  move- 
ments of  the  stomach  and  intestines.  As  compared  with  skeletal  and  cardiac 
muscle  it  is  a  much  more  slowly  acting  contractile  tissue.  Isolated  strips  of 
smooth  muscle,  as  a  rule,  contract  only  when  stimulated,  though  preparations 


FIG  333- — Contraction  Area  in  Smooth  Muscle.  A,  Showing  the  contraction  nodes 
of  the  fibers,  the  deep  staining  of  the  nodes,  the  condensation  of  surrounding  connective 
tissue;  B,  diagrammatic,  showing  the  thickening  of  the  longitudinal  fibrillae.  T— ••— »:  ~* 
dog.  (Caroline  McGiil.) 


Intestine  of 


of  certain  tissues,  like  the  stomach  muscle  of  the  frog,  give  off  rhythmic 
contractions  occasionally.  In  this  regard  smooth  muscle  stands  intermedi- 
ate between  skeletal  and  cardiac  muscle;  the  former  is  normally  never 
automatic,  the  latter  always. 

Smooth  muscle  requires  a  different  type  of  stimulus  to  produce  contrac- 
tion; the  stimulus  must  be  more  prolonged  and  more  intense.  For  example, 
smooth  muscle  is  not  readily  responsive  to  induction  currents  of  short  dura- 
tion, but  is  readily  stimulated  by  galvanic  currents  or  induction  currents  of 
longer  duration.  The  stimulus  must  be  applied  through  a  longer  interval 
of  time.  Preparations  of  the  stomach  muscle  can  scarcely  be  made  to 


CHANGES  DURING  THE  CONTRACTION  OF  SMOOTH  MUSCLE        531 

contract  by  a  single  induction  current,  no  matter  how  intense.  Such  muscle 
in  the  body  is  always  associated  with  the  local  nervous  apparatus  which 
plays  an  indeterminate  part  in  its  activity. 

The  ureters  and  gall-bladder  are  the  parts  most  difficult  to  excite  by 
stimuli;  they  do  not  act  at  all  till  the  stimulus  has  been  long  applied,  and  then 
contract  feebly  and  to  a  small  extent.  The  contractions  of  the  cecum  and 
stomach  are  quicker,  and  still  quicker  those  of  the  iris  and  of  the  urinary 
bladder.  The  contractions  of  the  small  and  large  intestines,  of  the  vas 
deferens,  and  of  the  pregnant  uterus  are  yet  more  regular  and  more 
sustained. 

Changes  During  the  Contraction  of  Smooth  Muscle. — The  dura- 
tion as  well  as  type  of  contraction  in  smooth  muscle  is  very  markedly  differ- 
ent from  that  of  voluntary  muscle.  A  contraction  in  smooth  muscle  is 


FIG.  334. — Showing  periodic  contractions  of  an  isolated  half  of  the  uterus  and 
Fallopian  tube  of  the  rat.  The  contractions  occur  spontaneously  in  serum  and  warm 
oxygenated  Ringer's  solution.  Experiment  by  Bonham.  Time  in  minutes. 

characterized  by  a  very  long  latent  period,  a  slowly  developed  contraction 
phase,  and  an  extremely  delayed  relaxation,  figure  356,  A.  The  amount  and 
duration  of  the  contractions  are  dependent  upon  the  strength  and  duration  of 
the  stimulus,  though  the  curve  of  contraction  itself  does  not  in  other  respects 
differ  sharply  from  the  type  of  curve  of  the  simple  muscle  contraction. 

Owing  to  the  apparently  different  structural  type  of  smooth  muscle,  es- 
pecial interest  attaches  to  the  changes  which  occur  during  its  contraction. 
Caroline  McGill  has  recently  re-examined  the  histological  structure  and  in- 
vestigated the  function  of  this  type  of  muscle,  and  we  are  able  to  present  a 
figure  showing  the  changes.  The  longitudinal  fibrillae,  which  are  readily 
stained  with  iron  hematoxylin,  show  distinct  shortening  and  thickening  at  the 


532  MUSCLE-NERVE    PHYSIOLOGY 

nodes  of  contraction  of  the  muscle,  figure  333,  B.  The  whole  fiber  is  thick- 
ened at  the  contraction  nodes  and  stains  very  readily  and  usually  uniformly. 
However,  by  certain  stains  the  fibrillae  can  be  traced  through  the  node.  The 
node  is  an  apparent  area  of  chemical  differentiation.  There  is  a  marked  con- 
densation of  the  intermuscular  fibrous  tissue,  which  is  doubtless  purely  a 
passive  phenomenon.  The  most  striking  change  during  contractions  is 
observed  in  the  nucleus,  figure  333,  A,  and  figure  334.  The  nucleus  during 
rest  is  a  long  slender  oval  or  spindle  with  a  general  chromatic  network. 
1  'During  contraction,  the  smooth  muscle  nuclei  shorten  and  thicken  by  an 
active  process.  The  chromatin  collects,  chiefly  at  the  two  ends  of  the  nu- 
cleus, leaving  a  relatively  clear  area  in  the  center." 

Ciliary  Motion. — Ciliary  motion,  which  is  closely  allied  to  ameboid 
and  muscular  motion,  is  alike  independent  of  the  will,  of  the  direct  influ- 
ence of  the  nervous  system,  and  of  muscular  contraction.  It  may  continue 
for  several  hours  after  death,  or  removal  of  the  ciliated  tissue,  provided  the 
portion  of  tissue  under  examination  be  kept  moist.  Its  independence  of  the 
nervous  system  is  shown  also  in  its  occurrence  in  the  lowest  invertebrate 
animals  which  are  apparently  unprovided  with  anything  analogous  to  a 
nervous  system,  and  in  its  persistence  when  the  ciliated  cells  are  completely 
separated  from  each  other  by  teasing  out  in  serum  or  other  physiological 
solution.  The  vapor  of  chloroform  arrests  the  motion;  but  it  is  renewed 
on  the  discontinuance  of  the  application  of  the  anesthetic.  The  movement 
ceases  when  the  cilia  are  deprived  of  oxygen  (although  it  may  continue  for 
a  time  in  the  absence  of  free  oxygen),  but  is  revived  on  the  admission  of  this 
gas.  Carbon  dioxide  also  stops  the  movement.  The  contact  of  various 
substances,  e.g.,  bile,  strong  acids,  and  alkalies,  will  stop  the  motion  alto- 
gether; but  this  depends  chiefly  on  destruction  of  the  delicate  substance  of 
which  the  cilia  are  composed.  Temperatures  above  45°  C.  and  below  o°  C. 
stop  the  movement,  whereas,  moderate  heat  and  faintly  alkaline  solutions 
are  favorable  to  the  action  and  revive  the  movement  after  temporary  cessation. 
The  exact  explanation  of  ciliary  movement  is  not  known.  Whatever  may 
be  the  exact  explanation,  the  movement  must  depend  upon  some  changes 
going  on  in  the  cells  of  which  the  cilia  are  a  part  and  not  on  changes  limited 
to  the  cilia  themselves,  since,  when  the  latter  are  cut  off  from  the  cell  the 
movement  ceases,  and  when  severed  so  that  portions  of  the  cilia  are  left 
attached  to  the  cell,  the  attached  and  not  the  severed  portions  continue  the 
movement.  Ciliary  contraction  is  to  be  regarded  as  a  type  of  motor  activity 
carried  out  in  a  specialized  form  of  motor  apparatus.  The  changes  going 
on  in  the  cell  must  be  classed  with  similar  changes  in  heart  or  skeletal 
muscle.  Ciliary  tissue  is  like  cardiac  in  at  least  two  characteristics:  the  cells 
are  capable  of  conducting  a  stimulus  from  cell  to  cell,  and  ciliary  activity 
is  automatic.  As  a  special  illustration  of  cilia-like  action  may  be  mentioned 
the  motion  of  spermatozoa,  which  are  cells  with  a  single  cilium. 


THE    FUNCTION    OF    NERVE    FIBER  533 


THE  FUNCTION  OF  NERVE  FIBER. 

The  Nerve  Impulse. — The  motor  nerve  fibers  of  the  muscle-nerve 
preparation  are  of  the  medullated  type  described  on  page  66.  But  the  es- 
sential structure,  possessed  by  all  fibers,  is  the  axis  cylinder.  The  peculiar 
function  of  the  nerve  fiber,  i.e  ,  of  the  axis  cylinder,  is  its  power  to  conduct 
a  physiological  change  along  its  extent,  a  phenomenon  known  as  a  nerve 
impulse.  A  normal  nerve  impulse  in  a  motor  nerve  has  its  origin  in  the 
motor  cell  of  the  central  nervous  system  of  which  the  fiber  is  an  outgrowth. 
The  manner  in  which  such  discharge  from  the  cell  takes  place  will  be  dis- 
cussed later.  But  nerve  impulses  may  be  aroused  by  various  artificial 
means,  they  are  influenced  by  certain  conditions  in  the  environment,  and 
possess  certain  other  properties  that  may  be  discussed  at  this  point. 

Nerve  Stimuli. — Nerve  fibers  like  skeletal  muscle  require  stimulation 
before  they  can  manifest  any  of  their  properties,  since  they  have  no  power 
of  themselves  of  originating  nerve  impulses.  The  stimuli  which  are  capable 
of  exciting  nerves  to  action  are,  as  in  the  case  of  muscle,  very  diverse.  The 
mechanical,  chemical,  thermal,  and  electrical  stimuli  which  may  be  used 
in  the  case  of  muscles  are  also,  with  certain  differences  in  the  methods  em- 
ployed, efficacious  in  stimulating  the  nerve.  The  chemical  stimuli  are  chiefly 
these:  withdrawal  of  water  as  by  drying;  strong  solutions  of  neutral  salts  of 
potassium,  sodium,  etc.;  free  inorganic  acids,  except  phosphoric;  and  some 
organic  acids.  The  electrical  stimuli  employed  are  the  induction  and  con- 
tinuous currents  concerning  which  the  observations  in  reference  to  muscular 
irritability  should  be  consulted.  Galvanic  currents  stimulate  nerves  only  at 
the  moment  of  turning  on  the  current  and  of  turning  it  off.  Weaker  electrical 
stimuli  will  excite  nerves  than  will  excite  muscles;  the  nerve  impulse  appears 
to  gain  strength  as  it  descends,  and  a  weaker  stimulus  applied  far  from  the 
muscle  will  have  the  same  effect  as  a  slightly  stronger  one  applied  to  the 
nerve  near  the  muscle. 

Characteristics  of  the  Nerve  Impulse. — When  a  nerve  impulse  is 
aroused  in  a  motor  nerve,  as  by  stimulating  a  nerve  in  its  course  by  an  in- 
duced current  of  medium  strength,  it  is  propagated  along  the  axis  cylinder 
to  the  muscle  where  it  arouses  a  contraction  of  the  muscle  fiber.  In  the  con- 
traction of  the  muscle  we  have  indirect  but  conclusive  evidence  of  the  passage 
of  the  nerve  impulse,  for  it  can  be  readily  proven  that  the  electrical  current 
does  not  escape  to  the  muscle.  In  this  instance  it  can  be  shown  that  there 
is  a  nerve  impulse  passing  from  the  point  of  stimulation  in  the  direction  away 
from  the  muscle;  i.e.j  the  artificially  aroused  nerve  impulse  passes  over  the 
entire  extent  of  the  fiber  stimulated.  In  fact,  a  nerve  impulse  is  known  to 
travel  from  its  point  of  origin  over  the  entire  neurone  affected.  This  anti- 
dromal  nerve  impulse,  of  course,  does  not  exist  in  the  normal  case,  since  the 


534  MUSCLE-NERVE    PHYSIOLOGY 

normal  nerve  impulse  arises  in  the  nerve-cell  body  and  passes  out  over  the 
fiber  from  its  origin  to  its  extremity. 

The  nerve  impulse  travels  over  the  nerve  fiber  with  a  velocity  that  was 
first  determined  by  Helmholtz.  He  found  that  in  the  sciatic  of  the  frog  the 
nerve  impulse  travels  at  the  rate  of  twenty-seven  meters  per  second.  The 
rate  has  been  measured  in  a  number  of  animals  and  varies  between  wide 
limits.  In  human  nerves  the  rate  is  variously  given,  but  Helmholtz'  figure 
of  thirty-four  meters  per  second  may  be  taken  as  a  fair  average. 

The  presence  of  the  nerve  impulse  can  be  detected  by  the  action  current, 
which  exists  in  nerve  as  in  muscle  (see  page  494  for  methods  of  detecting 
the  action  current). 

Rheoscopic  Frog. — The  action  current  may  be  demonstrated  by  means  of 
the  following  experiment: 

The  muscle  current  produced  by  stimulating  the  nerve  of  one  muscle-nerve 
preparation  may  be  used  to  stimulate  the  nerve  of  a  second  muscle-nerve 
preparation.  The  hind  leg  of  a  frog  with  the  nerve  going  to  the  gastrocnemius 
cut  long  is  placed  upon  a  glass  plate  and  arranged  in  such  way  that  its  nerve 
touches  in  two  places  the  gastrocnemius  muscle,  exposed  but  preserved  in  situ 
in  the  opposite  thigh  of  the  frog.  The  electrodes  from  an  induction  coil  are 
placed  behind  the  sciatic  nerve  of  the  second  preparation,  high  up.  On 
stimulating  it  with  a  single  induction  shock,  the  muscles  not  only  of  the  same 
leg  are  found  to  undergo  a  twitch,  but  also  those  of  the  first  preparation,  al- 
though this  is  not  near  the  electrodes.  The  stimulation  cannot  be  due  to  an 
escape  of  the  stimulating  current  into  the  first  nerve,  but  is  due  to  the  action 
current  of  the  second  muscle.  This  experiment  is  known  under  the  name  of 
the  rheosco pic  frog. 

When  the  nerve  impulse  is  studied  by  means  of  the  action  current  it  is 
found  that  a  nerve  impulse  can  be  aroused  by  a  weaker  stimulus  than  is  re- 
quired to  produce  a  minimal  contraction  of  a  muscle.  The  response  of  the 
nerve  to  graduated  strengths  of  the  stimulus  is  increased  very  rapidly  with 
slight  increase  of  strength  of  the  stimulus,  the  augmentation  extending 
through  a  somewhat  greater  range  than  for  muscle.  If  the  stimulus  is  still 
further  increased  there  is  only  slight  increase  of  the  resulting  nerve  impulse. 

Fatigue  of  Nerve  Fiber. — Many  efforts  have  been  made  to  discover 
evidences  of  fatigue  of  nerve  fiber,  with  practically  completely  negative 
results.  A  difficulty  has  been  to  secure  means  of  measuring  change  in  in- 
tensity of  the  nerve  impulse.  The  muscle  quickly  fatigues  so  that  the 
character  of  the  muscle  response  cannot  be  taken  when  measured  in  the 
ordinary  way.  An  effective  method  used  by  Howell,  Budgett,  and  Leonard 
consists  in  cooling  a  segment  of  nerve  to  suspend  its  conductivity,  during 
stimulation  of  the  free  end,  and  periodically  warming  up  the  cooled  segment 
of  nerve  to  test  the  strength  of  nerve  impulse  passing  through  it  to  the  un- 
fatigued  muscle  beyond.  By  this  and  other  methods  it  has  been  found  that 


THE    EFFECTS    OF    BATTERY    CURRENTS    ON    NERVE  FIBER         535 

a  motor  nerve  is  riot  fatigued  by  at  least  ten  hours'  continuous  stimulation 
with  induction  currents. 

One  must  hesitate  to  draw  the  conclusion,  however,  that  the  nerve  fiber 
conducts  the  nerve  impulse  without  loss  of  energy.  The  fiber  can  be 
anesthetized,  it  responds  to  temperature  changes,  and  gives  other  evidences 
of  susceptibility  to  conditions  which  influence  metabolism  in  other  forms  of 
protoplasm.  Perhaps  the  nerve  fiber  is  capable  of  repairing  its  wastes  as 
rapidly  as  they  occur. 

The  Effects  of  Battery  Currents  on  Nerve  Fiber. — Galvanic  currents 
influence  nerves  in  ways  that  call  for  special  discussion.  A  constant  cur- 
rent, say  from  a  Daniell  battery,  can  be  introduced  into  the  nerve  of  a  muscle- 
nerve  preparation  by  means  of  a  pair  of  non-polarizable  electrodes,  figure 
323,  and  a  convenient  key  arranged  for  turning  the  current  on  or  off  the 
nerve.  It  will  be  found  that  with  a  current  of  moderate  strength  there  will 
be  a  contraction  of  the  muscle,  both  at  the  closing  and  the  opening  of  the  key 
(called,  respectively,  making  and  breaking  contractions),  but  that  during  the 
interval  between  these  two  events  the  muscle  remains  flaccid,  provided  the 
battery  current  continues  of  constant  intensity.  If  the  current  be  a  very 
weak  or  a  very  strong  one,  the  effect  is  not  quite  the  same;  one  or  the  other 
of  the  contractions  may  be  absent.  Which  of  these  contractions  is  absent 
depends  upon  another  circumstance,  viz.,  the  direction  of  the  current.  The 
direction  of  the  current  may  be  ascending  or  descending:  if  ascending,  the 
anode  or  positive  pole  is  nearer  the  muscle  than  the  cathode  or  negative  pole, 
and  the  current  to  return  to  the  battery  has  to  pass  up  the  nerve;  if  descend- 
ing, the  position  of  the  electrodes  is  reversed.  It  will  be  necessary  before 
considering  this  question  further  to  return  to  the  apparent  want  of  effect  of 
the  constant  current  during  the  interval  between  the  make  and  the  break 
contraction.  To  all  appearances  no  change  is  produced,  but  in  reality  a  very 
important  alteration  of  the  irritability  and  conductivity  is  brought  about  in 
the  nerve  by  the  passage  of  this  constant  or  polarizing  current. 

A  second  way  of  showing -the  effect  of  the  polarizing  current  is  by  stimu- 
lating the  nerve  by  a  pair  of  electrodes  from  an  induction  coil,  while  the  polar- 
izing current  from  the  battery  is  flowing  through  the  nerve.  If  the  strength 
of  stimulus  required  in  order  that  a  minimum  contraction  be  obtained  by  the 
induction  shock  before  the  polarizing  current  is  applied  and  the  secondary 
coil  be  removed  slightly  further  from  the  primary,  the  induction  current 
cannot  now  produce  a  contraction.  If,  now,  the  polarizing  current  be  sent  in 
a  descending  direction,  that  is  to  say,  with  the  cathode  nearest  the  muscle,  and 
the  induction  current  which  was  before  insufficient  be  applied  between  the 
cathode  and  the  muscle;  it  will  now  prove  sufficient  to  cause  a  contraction. 
This  indicates  that  with  a  descending  current  the  irritability  of  the  nerve  is 
increased  at  the  cathode.  If,  instead  of  applying  the  induction  electrodes 
below  the  polarizing  electrodes,  they  are  applied  above  them,  the  irritability  of 


536 


MUSCLE-NERVE   PHYSIOLOGY 


the  nerve  is  found  to  be  decreased.  If  the  polarizing  current  is  reversed,  i.e., 
made  ascending,  then  the  condition  of  irritability  of  the  nerve  is  reversed. 
Both  methods  show  that  the  polarization  consists  in  an  increase  in  irritability 
at  the  cathode,  called  catelectrotonus,  and  a  decrease  at  the  anode  called 
anelectrotonus.  The  total  change  is  called  by  the  term  electrotonus.  As  there 
is  between  the  electrodes  both  an  increase  and  a  decrease  of  irritability  on  the 
passage  of  a  polarizing  current,  it  is  evident  that  there  must  be  a  neutral  point 
where  there  is  neither  increase  nor  decrease  of  irritability.  The  position  of 
this  neutral  point  is  found  to  vary  with  the  intensity  of  the  polarizing  current; 
when  the  current  is  weak  the  point  is  nearer  the  anode,  when  strong  nearer  the 


FlG-  335- — Diagram  Illustrating  the  Effects  of  Various  Intensities  of  the  Polarizing 
Currents,  n,  n',  Nerve;  a,  anode;  k,  cathode;  the  curves  above  indicate  increase,  and 
those  below  decrease  of  irritability,  and  when  the  current  is  small  the  increase  and  decrease 
are  both  small,  with  the  neutral  point  near  a,  and  so  on  as  the  current  is  increased  in 
strength. 

cathode,  figure  335.  When  a  constant  current  passes  into  a  nerve,  therefore, 
if  a  contraction  result,  it  may  be  assumed  that  it  is  due  to  the  increased  irri- 
tability produced  in  the  neighborhood  of  the  cathode,  but  the  breaking  con- 
traction must  be  produced  by  a  rise  in  irritability  from  a  lowered  state  to  the 
normal  in  the  neighborhood  of  the  anode. 

The  contractions  produced  in  the  muscle  of  a  muscle-nerve  preparation 
by  a  constant  current  have  been  arranged  in  a  table  which  is  known  as 
Pfluger's  Law  of  Contractions.  It  is  really  only  a  statement  as  to  when  a 
contraction  may  be  expected: 


Strength  of  current  used 

Descending  current 

Ascending  current 

Make 

Break 

Make 

Break 

Very  weak  

Yes 
Yes 
Yes 
Yes 

No 

No 
Yes 
No 

No 
Yes 
Yes 

No 

No 
No 
Yes 
Yes 

Weak  

Moderate  

Strong  

EFFECT    OF    BATTERY    CURRENTS    ON    DEEP-SEATED    NERVES     537 

During  the  passage  of  a  constant  current  through  a  nerve  and  immediately 
after  its  cessation,  there  is  a  change  in  the  conductivity  as  well  as  of  the  irri- 
tability of  the  nerve  at  the  anode  and  cathode,  respectively.  During  the  pas- 
sage of  the  current,  the  conductivity  is  increased  at  the  cathode  and  decreased 
at  the  anode.  After  the  passage  of  the  current,  the  effect  is  reversed.  With 
strong  currents  the  area  of  decreased  conductivity  may  be  sufficient  to  act  as  a 
block,  preventing  the  passage  of  impulses  over  it. 

The  foregoing  statements  concerning  the  changes  produced  in  a  nerve 
by  the  passage  of  a  constant  current  may  be  briefly  summarized  as  follows: 

I.  A  nerve  is  more  irritable  to  the  closing  of  a  constant  current  than  it  is  to 
the  opening  of  a  constant  current. 

II.  During  the  passage  of  the  current  through  the  nerve,  both  its  irritability 
and  conductivity  are  increased  at  the  cathode  and  decreased  at  the  anode. 

III.  After  the  passage  of  the  current,  the  irritability  and  conductivity  are 
both  decreased  at  the  cathode  and  increased  at  the  anode. 

The  Effect  of  Battery  Currents  on  Deep-seated  Nerves. — The  follow- 
ing account  is  condensed  from  Lombard  in  "An  American  Text-book  of 
Physiology." 

As  an  electric  current  cannot  be  applied  to  living  human  nerves  directly, 
it  is  applied  to  the  skin  along  the  course  of  the  nerve.  The  current  passes 

Skin 


FIG.  33 50. — Diagram  of  Skin  and  Subjacent  Nerve.  A,  the  positive  electrode  or 
phvsical  anode;  B,  the  negative  electrode  or  physical  cathode.  Signs,  -f-  physiological 
anodes;  signs  —  physiological  cathodes.  (After  Waller.) 

from  the  anode  or  positive  pole  through  the  skin,  and  spreads  out  in  the 
tissues  much  as  the  bristles  of  a  brush;  it  then  gradually  concentrates  and 
leaves  the  skin  at  the  cathode  or  negative  pole. 

In  addition  to  the  physical  anode  and  cathode  of  the  battery,  there  are 
what  are  called  physiological  anodes  and  cathodes.  There  is  a  physiological 
anode  at  every  point  where  the  current  enters  a  nerve,  and  a  physiological 
cathode  at  every  point  where  it  leaves  it. 

Generally  when  the  current  is  applied  to  nerves  through  the  skin,  only 
part  of  it  flows  longitudinally  along  the  nerves;  most  of  it  passes  diagonally 
across  them  to  the  tissues  below.  Thus  it  happens  that  in  that  part  of  the 
nerve  beneath  either  the  physical  anode  or  cathode,  groups  of  physiological 
anodes  and  cathodes  are  found. 

The  contraction  which  occurs  when  the  current  is  closed  (closing  con- 
traction) represents  irritation  at  the  physiological  cathode,  while  the  opening 


538 


MUSCLE-NERVE    PHYSIOLOGY 


contraction  represents  irritation  at  the  physiological  anode.  Since  there  are 
physiological  anodes  and  cathodes  beneath  each  electrode,  one  or  more  of 
four  conditions  may  arise : 

1.  Anodic  closing  contraction,  i..e.,  the  effect  of  the  change  developed  at 
the  physiological  cathode,  beneath  the  physical  anode  (positive  pole). 

2.  Anodic  opening  contraction,  ie.,  the  effect  of  the  change  developed  at 
the  physiological  anode,  beneath  the  physical  anode  (positive  pole). 

3.  Cathodic  closing  contraction,  i.e.,  the  effect  of  the  change  developed 
at  the  physiological  cathode,  beneath  the  physical  cathode  (negative  pole). 

4.  Cathodic  opening  contraction,  i.e.,  the  effect  of  the  change  developed 
at  the  physiological  anode,  beneath  the  physical  cathode  (negative  pole). 

The  following  abbreviations  of  these  contractions  are  used:  ACC,  AOC, 
KCC,  KOC. 

The  closing  contractions,  KCC  and  ACC,  are  stronger  than  the  opening 
contractions,  KOC  and  AOC.  Of  the  closing  contractions,  KCC  is  stronger 
than  ACC.  Of  the  opening  contractions,  AOC  is  stronger  than  KOC. 
These  facts  are  also  shown  in  a  table  of  the  effects  of  gradually  increasing  the 
strength  of  the  current. 

Weak  currents.  Medium  currents.  Strong  currents* 

KCC  KCC  KCC 

ACC  ACC 

AOC  AOC 

KOC 

Sometimes  AOC  is  stronger  than  ACC. 


r.  nervi  med.  m.  pron.  ? 
tereti. 


m.  palmaris  longus ~|-/ 


m.  ulnaris  int. 


n.  ulnaris 


/    \  —  -l m.  flex.  dig.  sublim. 


r.  vol.  prof.  n.  ulnar. 
m.  palmar  brevis 
m.  abduc.  dig.  min. 
m.  flex.  dig.  min. 
m.  oppon.  dig.  min.  -- 

mm. lumbr.  II.,III.,IV.  \-'-\  '--"-' 


m.  radial.  Intern. 

m.  flex.  dig.  prof. 


U—      --  m.  flex.  poll.  long. 
m.  medianus 

•-X- m.  abduc.  poll.  brev. 

L-.  _. .A.        m.  oppon.  poll. 

m.  flex.  poll.  brev. 

.  lumbric.  I. 


FIG.  336. — Figure  Showing  Motor  Points  in  the  Forearm. 


LOCOMOTION  539 

In  diseases  which  cause  degeneration  of  the  nerves  going  to  a  muscle, 
stimulation  causes  results  different  from  the  above,  and  we  get  what  is  known 
as  the  reaction  of  degeneration. 

The  intensity  of  the  anodic  or  cathodic  effects  is  increased  by  using  small 
electrodes,  and  decreased  by  electrodes  of  large  surface.  In  fact  in  practice 
it  is  usual  to  apply  the  indifferent  electrode  to  an  extended  surface,  thus  re- 
ducing its  effect  below  the  stimulating  intensity.  This  gives  only  one  active 
stimulating  electrode  and  is  known  as  the  method  of  unipolar  stimulation. 

SOME  SPECIAL  COORDINATED  MOTOR  ACTIVITIES. 

I.  LOCOMOTION. 

The  greater  number  of  the  more  important  muscular  actions  of  the  human 
body,  those,  namely,  which  are  arranged  harmoniously  so  as  to  subserve 
some  definite  purpose  in  the  animal  economy,  are  described  in  various  parts 
of  this  work  in  the  sections  which  treat  of  the  physiology  of  the  processes  by 
which  these  muscular  actions  are  resisted  or  carried  out.  There  are,  how- 
ever, some  very  important  and  somewhat  complicated  muscular  acts  which 
may  be  best  described  in  this  place. 

Walking. — The  coordinated  movements  of  the  body  are  carried  out 
by  the  skeletal  muscles  acting  on  the  skeletal  elements  as  a  system  of  levers. 
Even  the  bones  of  the  skull  are  levers  in  so  far  as  their  relations  to  muscles 
are  concerned. 

Examples  of  the  Three  Orders  of  Levers  in  the  Human  Body. — All  levers 
have  been  divided  into  three  kinds,  according  to  the  relative  position  of  the 
power,  the  weight  to  be  moved,  and  the  axis  of  motion  or  fulcrum.  In  a  lever  of 


f        f       Y 

j  !  O 

A 


FIG.  337. 

the  first  kind  the  power  is  at  one  extremity  of  the  lever,  the  weight  at  the 
other,  and  the  fulcrum  between  the  two.  If  the  initial  letters  only  of  the 
power,  weight,  andfulcmm  be  used,  the  arrangement  will  stand  thus:  P.  F.  W. 
A  poker  as  ordinarily  used,  or  the  bar  in  figure  337,  may  be  cited  as  an  example 
of  this  variety  of  lever;  while,  as  an  instance  in  which  the  bones  of  the  human 


540 


MUSCLE-NERVE   PHYSIOLOGY 


skeleton  are  used  as  a  lever  of  the  same  kind,  may  be  mentioned  the  act  of 
raising  the  body  from  the  stooping  posture  by  means  of  the  hamstring  muscles 
attached  to  the  tuberosity  of  the  ischium  or  of  the  triceps  which  extends  the 
forearm  by  action  at  the  elbow,  figure  337. 

In  a  lever  of  the  second  kind,  the  arrangement  is  thus:  P.  W.  F.;  and  this 
leverage  is  employed  in  the  act  of  raising  the  handles  of  a  wheelbarrow,  or  in 
stretching  an  elastic  band,  as  in  figure  338.  In  the  human  body  the  act  of 
opening  the  mouth  by  depressing  the  lower  jaw  is  an  example  of  the  same  kind 


FIG.  338. 

— the  tension  of  the  muscles  which  close  the  jaw  representing  the  weight, 
figure  338. 

In  a  lever  of  the  third  kind  the  arrangement  is  F.  P.  W.,  and  the  act  of 
raising  a  pole,  as  in  figure  339,  is  an  example.  In  the  human  body  there  are 
numerous  examples  of  the  employment  of  this  kind  of  leverage.  The  act  of 
bending  the  forearm  may  be  mentioned  as  an  instance,  figure  339.  The  act 
of  biting  is  another  example. 

At  the  ankle  we  have  examples  of  all  three  kinds  of  lever,  ist  kind — Ex- 
tending the  foot.  3d  kind — Flexing  the  foot.  In  both  these  cases  the  foot 


FIG.  339. 

represents  the  weight,  the  ankle  joint  the  fulcrum,  the  power  being  the  gas- 
trocnemius  muscles  in  the  first  case  and  the  tibialis  anticus  in  the  second 
case,  and  kind — When  the  body  is  raised  on  tiptoe.  Here  the  tip  of  the 
toe  is  the  fulcrum,  the  weight  of  the  body  acting  at  the  ankle  joint  the 
weight,  and  the  gastrocnemius  muscles  the  power. 

In  the  human  body,  levers  are  most  frequently  used  at  a  disadvantage  as 
regards  power,  the  latter  being  sacrificed  for  the  sake  of  a  greater  range  of 
motion.  Thus  in  the  diagrams  of  the  first  and  third  kinds  it  is  evident  that  the 
power  is  so  close  to  the  fulcrum  that  great  force  must  be  exercised  in  order  to 
produce  motion.  It  is  also  evident,  however,  from  the  same  diagrams,  that 


LOCOMOTION 


541 


by  the  closeness  of  the  power  to  the  fulcrum  a  great  range  of  movement  can 
be  obtained  by  means  of  a  comparatively  slight  shortening  of  the  muscular 
fibers. 

In  the  act  of  walking,  almost  every  voluntary  muscle  in  the  body  is  brought 
into  play,  either  directly  for  purposes  of  progression,  or  indirectly  for  the 
proper  balancing  of  the  head  and  trunk.  The  muscles  of  the  arms  are 
least  concerned;  but  even  these  are  for  the  most  part  instinctively  in  action 
to  some  extent. 

Among  the  chief  muscles  engaged  directly  in  the  act  of  walking  are  those 
of  the  calf,  which,  by  pulling  up  the  heel,  pull  up  also  the  astragalus,  and  with 
it,  of  course,  the  whole  body,  the  weight  of  which  is  transmitted  through  the 
tibia  to  this  bone,  figure  340.  When  starting  to  walk,  say  with  the  left  leg 
this  raising  of  the  body  is  not  entirely  dependent  on  the  muscles  of  the  left 


calf,  but  the  trunk  is  thrown  forward  in  such  a  way  that  it  would  fall  prostrate 
were  it  not  that  the  right  foot  is  brought  forward  and  planted  on  the  ground  to 
support  it.  Thus  the  muscles  of  the  left  calf  are  assisted  in  their  action  by 
those  muscles  on  the  front  of  the  trunk  and  legs  which,  by  their  contraction, 
pull  the  body  forward;  and,  of  course,  if  the  trunk  form  a  slanting  line,  with 
the  inclination  forward,  it  is  plain  that  when  the  heel  is  raised  by  the  calf 
muscles,  the  whole  body  will  be  raised,  and  pushed  obliquely  forward  and 
upward.  The  successive  acts  in  taking  the  first  step  in  walking  are  repre- 
sented in  figure  340,  i,  2,  3,  etc. 

Now  it  is  evident  that  by  the  time  the  body  has  assumed  the  position  No. 
3,  it  is  time  that  the  right  leg  should  be  brought  forward  to  support  it  and 
prevent  it  from  falling  prostrate.  This  advance  of  the  right  leg  is  effected 
partly  by  its  mechanically  swinging  forward,  pendulum-wise,  and  partly  by 
muscular  action;  the  muscules  used  being — i,  those  on  the  front  of  the 
thigh,  which  bend  the  thigh  forward  on  the  pelvis,  especially  the  rectus 
femoris,  with  the  psoas  and  the  iliacus;  2,  the  hamstring  muscles,  which 
slightly  bend  the  leg  on  the  thigh;  and,  3,  the  muscles  on  the  front  of  the  leg, 
which  raise  the  front  of  the  foot  and  toes,  and  so  prevent  the  latter  in 
swinging  forward  from  striking  the  ground. 

The  second  part  of  the  act  of  walking,  which  has  been  just  described,  is 
shown  in  the  diagram,  4,  figure  340. 


542  MUSCLE-NERVE    PHYSIOLOGY 

When  the  right  foot  has  reached  the  ground  the  action  of  the  left  leg  has 
not  ceased.  The  calf  muscles  of  the  latter  continue  to  act,  and,  by  pulling  up 
the  heel,  throw  the  body  still  more  forward  over  the  right  leg,  now  bearing 
nearly  the  whole  weight,  until  the  time  when  the  left  leg  should  again  swing 
forward,  and  the  left  foot  be  planted  on  the  ground  to  prevent  the  body 
from  falling  prostrate.  As  at  first,  while  the  calf  muscles  of  one  leg  and  foot 
are  preparing,  so  to  speak,  to  push  the  body  forward  and  upward  from 
behind  by  raising  the  heel,  the  muscles  on  the  front  of  the  trunk  and  the 
same  leg  (and  of  the  other  leg,  except  when  it  is  swinging  forward)  are 
helping  the  act  by  pulling  the  legs  and  trunk,  so  as  to  make  them  incline 
forward,  the  rotation  in  the  inclining  occurring  mainly  at  the  ankle  joint. 
Two  main  kinds  of  leverage  are,  therefore,  employed  in  the  act  of  walking, 
and  if  this  idea  be  firmly  grasped,  the  details  will  be  understood  with  com- 
parative ease.  One  kind  of  leverage  employed  in  walking  is  essentially 
the  same  with  that  employed  in  pulling  forward  the  pole,  as  in  figure  339. 
And  the  other,  less  exactly,  is  that  employed  in  raising  the  handles  of  a 
wheelbarrow.  Now,  supposing  the  lower  end  of  the  pole  to  be  placed  in 
the  barrow,  we  should  have  a  very  rough  and  inelegant,  but  not  altogether 
bad  representation  of  the  two  main  levers  employed  in  the  act  of  walk- 
ing. The  body  is  pulled  forward  by  the  muscles  in  front,  much  in  the 
same  way  that  the  pole  might  be  by  the  force  applied  at  p,  while  the  raising 
of  the  heel  and  pushing  forward  of  the  trunk  by  the  calf  muscles  are  roughly 
represented  on  raising,  the  handles  of  the  barrow.  The  manner  in  which 
these  actions  are  performed  alternately  by  each  leg,  so  that  one  after  the  other 
is  swung  forward  to  support  the  trunk,  which  is  at  the  same  time  pushed 
and  pulled  forward  by  the  muscles  of  the  other,  may  be  gathered  from  the 
previous  description. 

There  is  one  more  thing  to  be  especially  noticed  in  the  act  of  walking. 
Inasmuch  as  the  body  is  being  constantly  supported  and  balanced  on  each 
leg  alternately,  and  therefore  on  only  one  at  the  same  moment,  it  is  evident 
that  there  must  be  some  provision  made  for  throwing  the  center  of  gravity 
over  the  line  of  support  formed  by  the  bones  of  each  leg,  as,  in  its  turn,  it 
supports  the  weight  of  the  body.  This  may  be  done  in  various  ways,  and 
the  manner  in  which  it  is  effected  is  one  element  in  the  differences  which 
exist  in  the  walking  of  different  people.  Thus  it  may  be  done  by  ah  in- 
stinctive slight  rotation  of  the  pelvis  on  the  head  of  each  femur  in  turn,  in 
such  a  manner  that  the  center  of  gravity  of  the  body  shall  fall  over  the  foot 
of  this  side.  Thus  when  the  body  is  pushed  onward  and  upward  by  the 
raising,  say,  of  the  right  heel,  as  in  figure  340,  3,  the  pelvis  is  instinctively 
by  various  muscles  made  to  rotate  on  the  head  of  the  left  femur  at  the 
acetabulum,  to  the  left  side,  so  that  the  weight  may  fall  over  the  line  of 
support  formed  by  the  left  leg  at  the  time  that  the  right  leg  is  swinging 
forward,  and  leaving  all  the  work  of  support  to  fall  on  its  fellow.  Such  a 


LOCOMOTION  543 

"rocking"  movement  of  the  trunk  and  pelvis,  however,  is  accompanied 
by  a  movement  of  the  whole  trunk  and  leg  over  the  foot  which  is  being 
planted  on  the  ground,  figure  341,  the  action  being  accompanied  with  a 
compensatory  outward  movement  at  the  hip,  more  easily  appreciated  by 
looking  at  the  figure  (in  which  this  movement  is  shown  exaggerated)  than 
from  the  description.  ( 

Thus  the  body  in  walking  is  continually  rising  and  swaying  alternately 
from  one  side  to  the  other,  as  its  center  of  gravity  has  to  be  brought  alternately 
over  one  or  the  other  leg;  and  the  curvatures  of  the  spine  are  altered  in  cor- 


FIG.  341. 

respondence  with  the  varying  position  of  the  weight  which  it  has  to  support. 
The  extent  to  which  the  body  is  raised  or  swayed  differs  much  in  different 
people. 

In  walking,  one  foot  or  the  other  is  always  on  the  ground.  The  act  of 
leaping  or  jumping  consists  in  so  sudden  a  raising  of  the  heels  by  the  sharp 
and  strong  contraction  of  the  gastrocnemius  muscles  that  the  body  is 
jerked  off  the  ground.  At  the  same  time  the  effect  is  much  increased  by 
first  bending  the  thighs  on  the  pelvis,  and  the  legs  on  the  thighs,  and  then 
suddenly  straightening  out  the  angles  thus  formed.  The  share  which  this 
action  has  in  producing  the  effect  may  be  easily  known  by  attempting  to 
leap  in  the  upright  posture,  with  the  legs  quite  straight. 

Running. — Running  is  performed  by  a  series  of  rapid  low  jumps  pro- 
duced by  each  leg  alternately;  so  that,  during  each  complete  muscular  act 
concerned,  there  is  a  moment  when  both  feet  are  off  the  ground. 

In  all  these  cases,  however,  the  description  of  the  manner  in  which  any 
given  effect  is  produced,  can  give  but  a  very  imperfect  idea  of  the  infinite 


544  MUSCLE-NERVE   PHYSIOLOGY 

number  of  combined  and   harmoniously  arranged  muscular  contractions 
which  are  necessary  for  even  the  simplest  acts  of  locomotion. 

II.  THE  PRODUCTION  OF  THE  VOICE. 

Before  commencing  the  consideration  of  the  nervous  system  and  the 
special  senses  it  will  be  convenient  to  consider  first  speech,  the  production  of 
the  human  voice,  and  the  physiology  of  the  larynx  as  a  muscular  apparatus. 

The  Larynx. — In  nearly  all  air-breathing  vertebrate  animals  there 
are  arrangements  for  the  production  of  sound,  or  voice,  in  some  parts  of  the 
respiratory  apparatus.  In  many  animals,  the  sound  admits  of  being  variously 
modified  and  altered  during  and  after  its  production;  and,  in  man,  one  such 
modification  occurring  in  obedience  to  dictates  of  the  cerebrum,  is  speech. 

It  has  been  proven  by  observations  on  living  subjects,  by  means  of  the 
laryngoscope,  as  well  as  by  experiments  on  the  larynx  taken  from  the  dead 
body,  that  the  sound  of  the  human  voice  is  the  result  of  the  vibration  of  the 
inferior  laryngeal  ligaments,  or  the  true  vocal  cords  which  bound  the  glottis, 
caused  by  currents  of  expired  air  impelled  over  their  edges.  If  a  free  opening 
exist  in  the  trachea,  the  sound  of  the  voice  ceases,  but  it  returns  if  the  opening 
is  closed.  An  opening  into  the  air-passages  above  the  glottis,  on  the  con- 
trary, does  not  prevent  the  voice  being  produced.  By  forcing  a  current  of 
air  through  the  larynx  in  the  dead  subject,  clear  vocal  sounds  are  elicited, 
though  the  epiglottis,  the  upper  ligaments  of  the  larynx  or  false  vocal  cords, 
the  ventricles  between  the  upper  ligaments  and  the  inferior  ligaments,  and 
the  upper  part  of  the  arytenoid  cartilages,  be  all  removed.  But  the  true 
vocal  cords  must  remain  entire  with  their  points  of  attachment,  and  be  kept 
tense  and  so  approximated  that  the  fissure  of  the  glottis  may  be  narrow. 

The  vocal  ligaments  or  cords,  therefore,  are  regarded  as  the  proper  organs 
for  the  production  of  vocal  sounds.  The  modifications  of  these  sounds  are 
effected,  as  will  be  presently  explained,  by  other  parts",  viz.,  by  the  tongue, 
teeth,  lips,  etc.  The  structure  of  the  vocal  cords  is  adapted  to  enable  them  to 
vibrate  like  tense  membranes,  for  they  are  essentially  composed  of  elastic 
tissue;  and  they  are  so  attached  to  the  cartilaginous  parts  of  the  larynx  that 
their  position  and  tension  can  be  variously  altered  by  the  contraction  of  the 
muscles  which  act  on  these  parts. 

Thus  it  will  be  seen  that  the  larynx  is  the  organ  of  voice.  It  may  be  said 
to  consist  essentially  of  the  two  vocal  cords  and  the  various  cartilaginous, 
muscular,  and  other  apparatus  by  means  of  which  not  only  can  the  aperture 
of  the  larynx  (rima  glottidis)  be  closed  against  the  entrance  and  exit  of  air 
to  or  from  the  lungs,  but  also  by  means  of  which  the  cords  themselves  can  be 
stretched  or  relaxed,  brought  together  and  separated  in  accordance  with  the 
conditions  that  may  be  necessary  for  the  air  in  passing  over  them  to  set  them 
vibrating  to  produce  the  various  sounds.  Their  action  in  respiration  has 
been  already  referred  to. 


ANATOMY    OF    THE    LARYNX 


545 


Anatomy  of  the  Larynx. — The  principal  parts  entering  into  the  formation  of 
the  larynx,  figures  342  and  343,  are — the  thyroid  cartilage ;  the  cricoid  cartilage; 
the  two  arytenoid  cartilages ;  and  the.  two  true  vocal  cords.  The  epiglottis, 
figure  343,  has  but  little  to  do  with  the  voice,  and  is  chiefly  useful  in  pro- 
tecting the  upper  part  of  the  larynx  from  the  entrance  of  food  and  drink  in 


FIG.  342. — Cartilages  of  the  Larynx  Seen  from  the  Front,  i  to  4,  Thyroid  cartilage;  i, 
vertical  ridge  or  pomum  Adami;  2,  right  ala;  3,  superior,  and  4,  inferior  cornu  of  the  right 
side;  5,  6,  cricoid  cartilage;  5,  inside  of  the  posterior  part;  6,  anterior  narrow  part  of  the 
ring;  7,  arytenoid  cartilages.  X  f . 

deglutition.  The  false  vocal  cords  and  the  ventricle  of  the  larynx,  which  is  a 
space  between  the  false  and  the  true  cord  of  either  side,  need  be  only  referred  to. 
Cartilages. — a,  The  thyroid  cartilage,  figure  342,  i  to  4,  does  not  form  a 
complete  ring  around  the  larynx,  but  only  covers  the  front  portion,  b,  The 
cricoid  cartilage,  figure  342,  5,  6,  on  the  other  hand,  is  a  complete  ring;  the 


Lig.  Ary.  epiglott. 


Cart.  Wrisbergii. 
Cart.  Santorini. 

Cart,  aryten. 
Troc.  muscul. 

Lig.  crico-aryten. 

Dig.  cerato-crico.  post.  sup. 

Cornu.  infer. 

Lig.  carat-crico.  post.  inf. 


Cart,  tracheae 


Pars,  membran. 


FIG.  343. — The  Larynx  as  Seen  From  Behind  after  Removal  of  the  Muscles.  The  cartilages 
and  ligaments  only  remain.     (Stoerk.) 


back  part  of  the  ring  being  much  broader  than  the  front.  On  the  top  of  this 
broad  portion  of  the  cricoid  are,  c,  the  arytenoid  cartilages,  figure  342,  7,  the 
connection  between  the  cricoid  below  and  arytenoid  cartilages  above  being  a 
joint  with  synovial  membrane  and  ligaments,  the  latter  permitting  tolerably 
free  motion  between  them. 

Joints  and  Ligaments. — The  thyroid  cartilage  is  also  connected  with  the 

35 


S46  MUSCLE-NERVE   PHYSIOLOGY 

cricoid,  not  only  by  ligaments,  but  also  by  joints  with  synovial  membranes; 
the  lower  cornua  of  the  thyroid  clasping  the  cricoid  between  them,  yet  not 
so  tightly  but  that  the  thyroid  can  revolve,  within  a  certain  range,  around  an 
axis  passing  transversely  through  the  two  joints.  The  vocal  cords  are  attached 
behind  to  the  front  portion  of  the  base  of  the  arytenoid  cartilages,  and  in 
front  to  the  re-entering  angle  at  the  back  part  of  the  thyroid;  it  is  evident, 
therefore,  that  all  movements  of  either  of  these  cartilages  must  produce  an 
effect  on  them  of  some  kind  or  other.  Inasmuch,  too,  as  the  arytenoid  carti- 
lages rest  on  the  top  of  the  back  portion  of  the  cricoid  cartilage,  and  are 
connected  with  it  by  capsular  and  other  ligaments,  all  movements  of  the 
cricoid  cartilage  must  move  the  arytenoid  cartilages,  and  also  produce  an 
effect  on  the  vocal  cords. 

a  b 


FIG.  344. — The  Cartilages  and  Ligaments  of  the  Larynx,  Viewed  from  the  Front,  a, 
Epiglottis;  &,  hyoid  bone;  e,  cartilago  tritica;  d,  thyro-hyoid  membrane;  e,  superior  cornu  of 
thyroid  cartilage;  j,  thyroid  notch;  g,  pomum  Adami;  h,  crico-thyroid  membrane;  *,  inferior 
cornu  of  thyroid  cartilage;/,  cricoid  cartilage.  (Cunningham.) 

Intrinsic  Muscles. — The  intrinsic  muscles  of  the  larynx  are  so  connected 
with  the  laryngeal  cartilages  that  by  their  contraction  alterations  in  the  condi- 
tion of  the  vocal  cords  and  glottis  are  produced.  They  are  usually  divided 
into  four  classes  according  to  their  action,  viz.,  into  abductors,  adductors, 
sphincters  and  tensors.  The  abductors,  the  crico-arytenoidei,  widen  the 
glottis  by  separating  the  cords;  the  adductors,  consisting  of  the  thyro-ary- 
epiglottici,  the  arytenoideus  posticus  seu  transversus,  the  thyro-arytenoidei 
externi,  the  crico-arytenoidei  laterals,  and  the  thyro-arytenoidei  interni,  approxi- 
mate the  vocal  cords,  diminish  the  rima  glottidis,  and  act  generally  as  sphinc- 
ters and  supporters  of  the  glottis.  Finally,  the  tensors  of  the  cords  put  the 
cords  on  the  stretch,  with  or  without  elongating  them;  the  tensors  are  the 
crico-thyroidei  and  the  thyro-arytenoidei  interni. 

The  attachments  and  the  action  of  the  muscles  will  be  readily  understood 
from  the  following  table.  All  the  muscles  are  in  pairs  except  the  arytenoideus 
posticus. 


ANATOMY    OF    THE    LARYNX 


547 


TABLE  OF  THE  SEVERAL  GROUPS  OF  THE  INTRINSIC  MUSCLES  OF  THE  LARYNX  AND 

THEIR  ATTACHMENTS. 


Group 


Muscle 


Attachments 


Action 


I. 

Abductors. 


Crico-aryt- 
enoidei 
postici. 


II.  and  III. 

Adductors 

and 
Sphincters. 


In    three 
layers: 

a.       Outer 
layer, 
Thyro- 
aryepi- 
glottici. 


b.  Middle 
layer. 

i.    Aryte- 
noideus 
posticus. 

ii.  Thyro- 
arytenoi- 
dei  ex- 

terni. 


This  pair  of  muscles  arises,  on  either  side, 
from  the  posterior  surface  of  the  cor- 
responding half  of  the  cricoid  cartilage. 
From  this  depression  their  fibers  con- 
verge on  either  side  upward  and  out- 
ward to  be  inserted  into  the  outer  angle 
of  the  base  of  the  arytenoid  cartilages 
behind  the  cricoarytenoidei  laterales. 

A  pair  of  muscles.  Flat  and  narrow, 
which  arise  on  either  side  from  the  pro- 
cessus  muscularis  of  the  arytenoid  car- 
tilage, then  passing  upward  and  inward 
cross  one  another  in  the  middle  line  to 
be  inserted  into  the  upper  half  of  the 
lateral  border  of  the  opposite  arytenoid 
cartilage  and  the  posterior  border  of  the 
cartilage  of  Santorini.  The  lower  fibers 
run  forward  and  downward  to  be  in- 
serted into  the  thyroid  cartilage  near 
the  commissure.  The  fibers  attached 
to  the  cartilage  of  Santorini  are  con- 
tinued forward  and  upward  into  the 
aryepiglottic  fold. 

A  single  muscle.  Half-quadrilateral, 
attached  to  the  borders  of  the  arytenoid 
cartilages,  its  fibers  running  horizontally 
between  the  two. 


A  pair  of  muscles.  Each  of  which  con- 
sists of  three  chief  portions.  The 
lower  and  principal  fibers  arise  from  thej 
lower  half  of  the  internal  surface  of  the 
thyroid  cartilage,  close  to  the  angle,  and 
from  the  fibrous  expansion  of  the  crico- 
thyroid  ligament,  and  are  inserted  into 
the  lateral  border  of  the  arytenoid  car- 
tilage. The  inner  fibers  to  the  lower 
half  of  this  border,  and  the  outer  fibers 
into  the  upper  half,  some  pass  to  the 
cartilage  of  Wrisberg  and  the  ary- 
epiglottic fold. 


Draw  inward  and 
backward  the  outer 
angle  of  arytenoid 
cartilages,  and  so 
rotate  outward  the 
processus  vocalis  and 
widen  the  glottis. 


Help  to  narrow  or 
close  the  rima 
glottidis. 


Draws  together  the 
arytenoid  cartilages 
and  also  depresses 
them.  When  the 
muscle  is  paralyzed, 
the  inter-cartilagin- 
ous part  of  the  cords 
cannot  come  to- 
gether. 


548 


MUSCLE-NERVE    PHYSIOLOGY 


TABLE  OF  THE  SEVERAL  GROUPS  OF  THE  INTRINSIC  MUSCLES  OF  THE  LARYNX  AND 
THEIR  ATTACHMENTS. — Continued. 


Group 

Muscle 

Attachments 

Action 

iii.    Crico- 

A  pair  of  muscles.     They  arise  on  either 

Approximate  the  vocal 

arytenoi- 

side  from  the  middle  third  of  the  upper 

cords  by  drawing  the 

dei  later- 

border  of  the  cricoid  cartilage  and  are 

processus  muscularis 

ales. 

inserted  into  the  whole  anterior  margin 

of  the  arytenoid  car- 

of the  base  of  the  arytenoid  cartilage. 

tilages  forward  and 

Some  of  their  fibers  join  the  thyroid- 

downward     and     so 

aryepiglottici. 

rotate  the  processus 

vocalis  inward. 

c.      Inner- 

A pair  of  muscles.     They  arise  on  either 

Render  the  vocal  cords 

most  layer, 

side,  internally  from  the  angle  of  the 

tense  and  rotate  the 

Thyro- 

thyroid  cartilage,  internal  to  the  last      arytenoid    cartilages 

arytenoi- 

described  muscle,  b.  iii.,  and,  running      and  approximate  the 

dei    in- 

parallel  to  and  in  the  substance  of  the 

processus  vocalis. 

terni. 

vocal  cords,  are  attached  posteriorly  to 

the  processus  vocalis  and  to  the  outer 

surface  of  the  arytenoid  cartilages. 

IV. 

Tensors. 

Crico-thy- 

A  pair  of  fan-shaped  muscles  attached  on 

The  thyroid  cartilage 

roidei. 

either  side  to  the  cricoid  cartilage  below;      being    fixed    by    its 

from  the  mesial  line  in  front  for  nearly      extrinsic  muscles,  the 

one-half   of   its   lateral    circumference;     front  of  the  cricoid 

« 

backward  the  fibers  pass  upward  and 

cartilage    is     drawn 

outward  to  be  attached  to  the  lower 

upward,  and  its  back, 

border  of  the  thyroid  cartilage  and  to 

with   the   arytenoids 

the  front  border  of  its  lower  cornea. 

attached,    is    drawn 

down.     Hence      the 

Thyro-ary- 

The  most  posterior  part  is  almost  a  dis- 

vocal cords  are  elon- 

tenoidei 

tinct  muscle  and  its  fibers  are  all  but 

gated     antero-poste- 

interni. 

horizontal:    sometimes   this   muscle   is 

riorly  and  put  upon 

described  as  consisting  of  two  layers, 

the    stretch.     Paral- 

superficial   with    cortical    fibers,    deep 

ysis  of  these  muscles 

with   oblique    fibers,    described    under 

causes    an    inability 

Group  III. 

to      produce      high 

notes. 

Nerve  Supply. — The  sensory  filaments  of  the  superior  laryngeal  branch  of 
the  vagus  supply  the  epithelial  lining  of  the  larynx,  giving  it  that  acute  sensi- 
bility by  which  the  glottis  is  guarded  against  the  ingress  of  foreign  bodies, 
or  of  irrespirable  gases.  The  contact  of  these  stimulates  the  nerve  endings; 
and  the  sensory  nerve  impulse  conveyed  to  the  medulla  oblongata,  whether 
accompanied  by  sensation  or  not,  arouses  motor  impulses  through  the  fila- 
ments of  the  recurrent  or  inferior  laryngeal  branch,  which  excite  contraction  of 


ANATOMY    OF    THE    LARYNX 


549 


the  muscles  that  close  the  glottis.  Both  these  branches  of  the  vagi  cooperate 
also  in  the  production  and  regulation  of  the  voice.  The  inferior  laryngeal 
determines  the  degree  of  contraction  of  the  muscles  that  vary  the  tension  of 
the  vocal  cords,  and  the  superior  laryngeal  conveys  to  the  brain  the  sensation 
which  indicates  the  state  of  contraction  of  these  muscles.  Both  the  branches 
cooperate  also  in  the  actions  of  the  larynx  in  the  ordinary  slight  dilatation  and 
contraction  of  the  glottis  in  the  acts  of  expiration  and  inspiration,  more 
evidently  in  the  acts  of  coughing  and  other  forcible  respiratory  movements. 

The  Laryngoscope. — This  is  an  instrument  employed  in  investigating  the 
condition  of  the  pharynx,  larynx,  and  trachea.  It  consists  of  a  large  concave 
mirror  with  perforated  center  and  of  a  smaller  mirror  fixed  in  a  long  handle. 
In  use  the  patient  is  placed  in  a  chair,  a  good  light  (argand  burner,  or  lamp) 
is  arranged  on  one  side  of,  and  a  little  above  his  head.  The  operator  fixes  the 


Lig.  ary-epiglott. 

Cart.  Wrisbergii. 
Cart.  Santorini. 

ram.  Aryten.  obliqu. 

m.  Crico-arytenoid  post. 

Cornu  inferior. 

Lig.  cerato-cric. 

Pars.  post,  inf.membrain. 
Pars,  cartilag. 


FIG.  345. — The  Larynx  as  Seen  from  Behind.     To  show  the  intrinsic  muscles  posteriorly. 

(Stoerk.) 

concave  mirror  round  his  head  in  such  a  manner  that  he  looks  through  the 
central  aperture  with  one  eye.  He  then  seats  himself  opposite  the  patient, 
and  so  adjusts  the  position  of  the  mirror,  which  is  for  this  purpose  provided 
with  a  ball  and  socket  joint,  that  a  beam  of  light  is  reflected  on  the  lips  of  the 
patient. 

The  patient  is  now  directed  to  throw  his  head  slightly  backward,  and  to 
open  his  mouth ;  the  reflection  from  the  mirror  lights  up  the  cavity  of  the  mouth, 
and  by  a  little  alteration  of  the  distance  between  the  operator  and  the  patient 
the  point  at  which  the  greatest  amount  of  light  is  reflected  by  the  mirror — in 
other  words  its  focal  length — is  readily  discovered.  The  small  mirror  fixed 
in  the  handle  is  then  warmed,  either  by  holding  it  over  the  lamp,  or  by  putting 
it  into  a  vessel  of  warm  water ;  this  is  necessary  to  prevent  the  condensation  of 
breath  upon  its  surface.  The  degree  of  heat  is  regulated  by  applying  the  back 
of  the  mirror  to  the  hand  or  cheek,  r/hen  it  should  feel  warm  without  being 
painful. 

After  these  preliminaries  the  patient  is  directed  to  put  out  his  tongue, 
which  is  held  by  the  left  hand  of  the  operator  gently  but  firmly  against  the 


550 


MUSCLE-NERVE    PHYSIOLOGY 


lower  teeth  by  means  of  a  handkerchief.  The  warm  mirror  is  passed  to  the 
back  of  the  mouth,  until  it  rests  upon  and  slightly  raises  the  base  of  the  uvula, 
and  at  the  same  time  the  light  is  directed  upon  it:  an  inverted  image  of  the 


FIG.  346.— The  Parts  of  the  Laryngoscope. 

larynx  and  trachea  will  be  seen  in  the  mirror.  If  the  dorsum  of  the  tongue 
be  alone  seen,  the  handle  of  the  mirror  must  be  slightly  lowered  until  the 
larynx  comes  into  view;  care  should  be  taken,  however,  not  to  move  the  mirror 


FIG.  347. — To  Show  the  Position  of  the  Operator  and  Patient  when  Using  the 

Laryngoscope. 

upon  the  uvula,  as  it  excites  retching.     The  observation  should  not  be  pro- 
longed, but  should  rather  be  repeated  at  short  intervals. 

The  structures  seen  will  vary  somewhat  according  to  the  condition  of  the 
parts  as  to  inspiration,  expiration,  phonation,  etc.     They  are  the  following: 


MOVEMENTS    OF    THE    VOCAL    CORDS  551 

first,  and  apparently  at  the  posterior  part,  the  base  of  the  tongue^  immediately 
below  which  is  the  accurate  outline  of  the  epiglottis,  with  its  cushion  or  tubercle, 
figure  348.  Then  are  seen  in  the  central  line  the  true  vocal  cords,  white  and 
shining  in  their  normal  condition.  In  the  inverted  image  the  cords  are  closer 
together  posteriorly.  Between  them  is  left  an  open  slit,  narrow  while  a  high 
note  is  being  sounded,  wide  during  a  deep  inspiration.  On  each  side  of  the 
true  vocal  cords,  and  on  a  higher  level,  are  the  false  vocal  cords.  Still  more 
externally  than  the  false  vocal  cords  is  the  aryteno-epiglottidean  fold,  in  which 
are  situated  upon  each  side  three  small  elevations ;  of  these  the  most  external 
is  the  cartilage  of  Wrisberg,  the  intermediate  is  the  cartilage  of  Santorini,  while 
in  front  and  somewhat  below  the  preceding  is  the  summit  of  the  arytenoid 
cartilage  seen  only  during  deep  inspiration.  The  rings  of  the  trachea,  and  even 
the  bifurcation  of  the  trachea  itself,  if  the  patient  be  directed  to  draw  a  deep 
breath,  may  be  occasionally  seen. 

Movements  of  the  Vocal  Cords. — The  position  of  the  vocal  cords  in  ordi- 
nary tranquil  breathing  is  so  adapted  by  the  muscles  that  the  opening  of  the 
glottis  is  wide  and  triangular,  figure  .348,  B,  becoming  a  little  wider  at  each 
inspiration,  and  a  little  narrower  at  each  expiration.  On  making  a  rapid 
and  deep  inspiration  the  opening  of  the  glottis  is  widely  dilated,  figure  348,  C, 
and  somewhat  lozenge-shaped. 

In  Vocalization. — At  the  moment  of  the  emission  of  a  note  the  opening  is 
narrowed,  the  margins  of  the  arytenoid  cartilages  being  brought  into  contact 
and  the  edges  of  the  vocal  cords  approximated  and  made  parallel  at  the  same 
time  that  their  tension  is  much  increased.  The  higher  the  note  produced,  the 
tenser  do  the  cords  become,  figure  348,  A;  and  the  range  of  a  voice  depends, 
of  course,  in  the  main,  on  the  extent  to  which  the  degree  of  tension  of  the 
vocal  cords  can  be  thus  altered.  In  the  production  of  a  high  note  the  vocal 
cords  are  brought  well  within  sight,  so  as  to  be  plainly  visible  with  the  help 
of  the  laryngoscope.  In  the  utterance  of  low  tones,  on  the  other  hand,  the 
epiglottis  is  depressed  and  brought  over  the  vocal  cords,  figure  349.  The 
epiglottis,  by  being  somewhat  pressed  down  so  as  to  cover  the  superior  cavity 
of  the  larynx,  serves  to  render  the  notes  deeper  in  tone  and  at  the  same  time 
somewhat  duller,  just  as  covering  the  end  of  a  short  tube  placed  in  front  of 
caoutchouc  tongues  lowers  the  tone.  In  no  other  respect  does  the  epiglottis 
appear  to  have  any  effect  in  modifying  the  vocal  sounds. 

The  degree  of  approximation  of  the  vocal  cords  also  usually  corresponds 
with  the  height  of  the  note  produced;  but  probably  not  always,  for  the  width 
of  the  aperture  has  no  essential  influence  on  the  pitch  of  the  note,  as  long  as 
the  vocal  cords  have  the  same  tension;  only  with  a  wide  aperture  the  tone  is 
more  difficult  to  produce  and  is  less  perfect,  the  rushing  of  the  air  through  the 
aperture  being  heard  at  the  same  time. 

No  true  vocal  sound  is  produced  at  the  posterior  part  of  the  aperture 
of  the  glottis,  the  part  of  the  aperture  which  is  formed  by  the  space  between 
the  arytenoid  cartilages.  For  if  the  arytenoid  cartilages  be  approximated  in 
such  a  manner  that  their  anterior  processes  touch  each  other,  but  yet  leave  an 


552 


MUSCLE-NERVE    PHYSIOLOGY 


opening  behind  them  as  well  as  in  front,  no  second  vocal  tone  is  produced  by 
the  passage  of  the  air  through  the  posterior  opening,  but  merely  a  rustling 
sound.  The  pitch  of  the  note  produced  is  the  same  whether  the  posterior 
part  of  the  glottis  be  open  or  not. 


FIG.  348. — Three  Laryngoscopic  Views  of  the  Superior  Aperture  of  the  Larynx  and 
Surrounding  Parts.  A,  The  glottis  during  the  emission  of  a  high  note  in  singing;  B,  in 
easy  and  quiet  inhalation  of  air;  C,  in  the  state  of  the  widest  possible  dilatation,  as  in 
inhaling  a  very  deep  breath.  The  diagrams  A',  B',  and  C',  show  in  horizontal  sections  of 
the  glottis  the  position  of  the  vocal  ligaments  and  arytenoid  cartilages  in  the  three  several 
states  represented  in  the  other  figures.  In  all  the  figures,  so  far  as  marked,  the  letters 
indicate  the  parts  as  follows,  viz.:  /,  the  base  of  the  tongue;  e,  the  upper  free  part  of  the 
epiglottis;  e',  the  tubercle  or  cushion  of  the  epiglottis;  ph,  part  of  the  anterior  wall  of  the 
pharynx  behind  the  larynx;  in  the  margin  of  the  aryteno-epiglottidean  fold  w,  the  swelling 
of  the  membrane  caused  by  the  cartilages  of  Wrisberg;  s,  that  of  the  cartilages  of  Santorini; 
a,  the  tip  or  summit  of  the  arytenoid  cartilages;  cv,  the  true  vocal  cords  or  lips  of  the  rima 
glottidis;  cvs,  the  superior  or  false  vocal  cords;  between  them  the  ventricle  of  the  larynx; 
in  C,  tr  is  placed  on  the  anterior  wall  of  the  receding  trachea,  and  b  indicates  the  com- 
mencement of  the  two  bronchi  beyond  the  bifurcation  which  may  be  brought  into  view 
in  this  state  of  extreme  dilatation.  (Quain,  after  Czermak.) 

The  Voice  in  Singing. — The  laryngeal  tones  may  be  produced  in  three 
different  kinds  of  sequence.  The  first  is  the  monotonous,  in  which  the  notes 
have  nearly  all  the  same  pitch  as  in  ordinary  speaking;  the  variety  of  the 
sounds  of  speech  being  due  to  articulation  in  the  mouth.  In  speaking,  occa- 
sional syllables  receive  a  higher  intonation  for  the  sake  of  accent.  The 
second  mode  of  sequence  is  the  successive  transition  from  high  to  low  notes, 


THE    VOCAL    RANGE    OF    THE    VOICE  553 

and  vice  versa,  without  intervals;  such  as  is  heard  in  the  crying  in  children 
and  in  the  howling  and  whining  of  dogs.  The  third  mode  of  sequence  of  the 
vocal  sounds  is  the  musical,  in  which  each  sound  has  a  determinate  number 
of  vibrations,  and  the  numbers  of  the  vibrations  in  the  successive  sounds  have 
the  same  relative  proportions  that  characterize  the  notes  of  the  musical  scale. 
The  different  sounds  made  by  the  musical  voice  are  characterized  by  the 
three  properties  of  tones  in  general,  viz.,  the  pitch ,  which  is  dependent  on  the 
rate  of  vibration  of  the  vocal  cords;  the  loudness,  which  depends  on  the  force  of 


FIG.  349. — View  of  the  Upper  Part  of  the  Larynx  as  Seen  by  Means  of  the  Laryngo- 
scope during  the  utterance  of  a  grave  note,  c,  Epiglottis;  s,  tubercles  of  the  cartilages  of 
Santorini;  a,  arytenoid  cartilages;  z,  base  of  the  tongue;  ph,  the  posterior  wall  of  the  pharynx. 
(Czermak.) 

the  vibration,  and  the  quality  or  timber,  which  is  dependent  on  the  resonance 
of  the  cavities  of  the  respiratory  apparatus,  particularly  of  the  mouth,  phar- 
ynx, and  nasal  cavities. 

The  Vocal  Range  of  the  Voice. — In  different  individuals  this  com- 
prehends one,  two,  or  three  octaves.  In  singers,  that  is,  in  persons  trained  in 
singing,  it  extends  to  three  or  more  octaves.  But  the  male  and  female  voices 
commence  and  end  at  different  points  of  the  musical  scale.  The  lowest  note 
of  the  female  voice  is  about  an  octave  higher  than  the  lowest  of  the  male  voice; 
the  highest  note  of  the  female  voice  about  an  octave  higher  than  the  highest  of 
the  male.  The  entire  scale  of  the  average  human  voice  includes,  from  the 
lowest  male  note  to  the  highest  female,  about  three  to  three  and  a  half  octaves. 
Some  remarkable  musical  voices  have  had  a  range  of  three  and  a  half  octaves. 
A  principal  difference  between  the  male  and  female  voice  is,  therefore,  in  their 
pitch.  But  they  are  also  distinguished  by  the  quality  of  the  tone.  The  voices 
of  men  and  of  women  differ  among  themselves,  both  in  the  general  pitch  and 
in  the  quality.  There  are  two  kinds  of  male  voices,  technically  called  the  bass 
and  tenor,  and  two  of  female  voices,  the  contralto  and  soprano,  all  differing 
from  each  other  in  general  pitch.  The  bass  voice  reaches  lower  than  the 
tenor,  and  its  strength  lies  in  the  low  notes.  The  contralto  voice  is  of  lower 
range  than  the  soprano,  and  is  strongest  in  the  lower  notes  of  the  female  voice. 
The  barytone  and  mezzo-soprano  voices  are  intermediate  in  range;  the  bary- 
tone being  intermediate  between  bass  and  tenor,  the  mezzo-soprano  between 
the  contralto  and  soprano.  The  difference  in  the  pitch  of  the  male  and  the 
female  voices  depends  primarily  on  the  different  size  of  the  larynx  and  the 


554  MUSCLE-NERVE   PHYSIOLOGY 

length  of  the  vocal  cords  in  the  two  sexes;  their  relative  lengths  in  men  and 
women  are  as  three  to  two. 

The  boy's  larynx  resembles  the  female  larynx.  His  vocal  cords  before 
puberty  are  not  two- thirds  the  length  of  the  adult  cords;  and  the  angle  of  the 
thyroid  cartilage  is  as  little  prominent  as  in  the  female  larynx.  Boy's  voices 
are  alto  and  soprano,  resembling  in  pitch  those  of  women,  but  louder,  and 
differing  somewhat  from  them  in  tone.  But,  after  the  larynx  has  undergone 
the  change  produced  during  the  period  of  development  at  puberty,  the  boy's 
voice  becomes  bass  or  tenor.  While  the  change  of  form  is  taking  place  the 
voice  becomes  imperfect,  frequently  hoarse  and  crowing,  and  is  unfitted  for 
singing  until  the  readjustment  of  the  larynx  is  complete  and  the  muscles 
which  control  the  vocal  cords  are  again  coordinated.  In  eunuchs  who  have 
been  deprived  of  the  testes  before  puberty,  the  voice  does  not  undergo  this 
change.  The  voice  of  most  old  people  is  deficient  in  tone,  unsteady,  and 
more  restricted  in  extent.  The  first  defect  is  owing  to  the  ossification  of  the 
cartilages  of  the  larynx  and  the  altered  condition  of  the  vocal  cords;  the  want 
of  steadiness  arises  from  the  loss  of  nervous  power  and  command  over  the 
muscles,  the  result  of  which  is  here,  as  in  other  parts,  a  tremulous  movement. 
These  two  causes  combined  render  the  voices  of  old  people  void  of  tone,  un- 
steady, and  weak. 

Most  persons  have  the  power,  if  at  all  capable  of  singing,  of  modulating 
their  voices  through  a  double  series  of  notes  of  different  character:  namely, 
the  notes  of  the  natural  voice,  or  chest-notes,  and  the  falsetto  notes.  The 
natural  voice,  which  alone  has  been  hitherto  considered,  is  fuller,  and  excites 
a  distinct  sensation  of  much  stronger  vibration  and  resonance  than  the 
falsetto  voice,  which  has  more  of  a  flute-like  character. 

The  Quality  of  the  Voice. — The  difference  in  quality  of  voices,  seen 
when  two  or  more  persons  sound  the  same  note,  is  due  to  differences  in 
resonance  in  the  cavities  of  the  mouth  and  larynx  and  also  of  the  nose.  The 
shape  of  the  roof  of  the  mouth,  the  regularity  of  the  teeth,  and  the  size  of  the 
tongue,  and  the  size  and  clearness  of  the  nasopharynx  are  all  factors.  The 
size  and  shape  of  the  larynx  and  mouth  cavity  which  influence  the  voice 
quality  can  be  controlled  to  some>  extent  during  singing,  and  this  is  a  special 
point  of  training  in  voice  culture. 

Speech. — Besides  the  musical  tones  formed  in  the  larynx  a  great  number 
of  other  sounds  can  be  produced  in  the  vocal  tubes,  between  the  glottis 
and  the  external  apertures  of  the  air-passages,  the  combination  of  which 
sounds  into  different  groups  to  designate  objects,  properties,  actions,  etc., 
constitutes  language.  The  languages  do  not  employ  all  the  sounds  which 
can  be  produced  in  this  manner,  the  combination  between  certain  sounds 
being  often  difficult.  Those  sounds  which  are  easy  of  combination  enter,  for 
the  most  part,  into  the  formation  of  the  greater  number  of  languages.  Each 
language  contains  a  certain  number  of  such  sounds,  but  in  no  one  are  all 


ARTICULATE    SOUNDS 


555 


brought  together.  On  the  contrary,  different  languages  are  characterized  by 
the  prevalence  in  them  of  certain  classes  of  these  sounds,  while  other  sounds 
are  less  frequent  or  altogether  absent. 

Articulate  Sounds. — The  sounds  produced  in  speech,  or  the  articu- 
late sounds,  are  commonly  divided  into  vowels  and  consonants:  the  distinc- 
tion between  which  is  that  the  sounds  for  the  former  are  generated  by  the 
larynx,  while  those  for  the  latter  are  produced  by  interruption  of  the  current 
of  air  in  some  part  of  the  air-passages  above  the  larynx.  The  term  consonant 
has  been  given  to  these  because  several  of  them  are  not  properly  sounded,  ex- 
cept, consonantly  with  a  vowel.  Thus,  if  it  be  attempted  to  pronounce  aloud 
the  consonants  b,  d,  and  g,  or  their  modifications,  p,  t,  k,  the  intonation  fol- 
lows them  only  in  their  combination  with  a  vowel.  To  recognize  the  essential 
properties  of  the  articulate  sounds,  it  is  necessary  first  to  examine  them  as  they 
are  produced  in  whispering,  and  then  investigate  which  of  them  can  also  be 
uttered  in  a  modified  character  conjoined  with  vocal  tone.  By  this  procedure 
we  find  two  series  of  sounds:  in  one  the  sounds  are  mute,  and  cannot  be 
uttered  with  a  vocal  tone;  the  sounds  of  the  other  series  can  be  formed  inde- 
pendently'of  voice,  but  are  also  capable  of  being  uttered  in  conjunction 
with  it. 

All  the  vowels  can  be  expressed  in  a  whisper  without  vocal  tone,  that  is, 
mutely.  These  mute  vowel  sounds  differ,  however,  in  some  measure,  as  to 
their  mode  of  production,  from  the  consonants.  All  the  mute  consonants  are 
formed  in  the  vocal  tube  above  the  glottis,  or  in  the  cavity  of  the  mouth  or 
nose,  by  the  mere  rushing  of  the  air  between  the  surfaces  differently  modified 
in  disposition.  But  the  sound  of  the  vowels,  even  when  mute,  has  its  source 
in  the  glottis,  though  its  vocal  cords  are  not  thrown  into  the  vibrations  neces- 
sary for  the  production  of  voice;  and  the  sound  seems  to  be  produced  by  the 
passage  of  the  current  of  air  between  the  relaxed  vocal  cords.  The  same 
vowel  sound  can  be  produced  in  the  larynx  when  the  mouth  is  closed,  the 
nostrils  being  open,  and  the  utterance  of  all  vocal  tone  avoided.  The  sound 
when  the  mouth  is  open,  is  so  modified  by  varied  forms  of  the  oral  cavity  as  to 
assume  the  characters  of  the  vowels  a,  e,  i,  o,  u,  in  all  their  modifications. 

The  cavity  of  the  mouth  assumes  the  same  form  for  the  articulation  of 
each  of  the  mute  vowels  as  for  the  corresponding  vowel  when  vocalized;  the 
only  difference  in  the  two  cases  lies  in  the  kind  of  sound  emitted  by  the 
larynx.  It  has  been  pointed  out  that  the  conditions  necessary  for  changing 
one  and  the  same  sound  into  the  different  vowels  are  differences  in  the  size  of 
two  parts — the  oral  canal  and  the  oral  opening;  and  the  same  is  the  case 
with  regard  to  the  mute  vowels.  By  oral  canal  is  meant  here  the  space 
between  the  tongue  and  palate:  for  the  pronunciation  of  certain  vowels 
both  the  opening  of  the  mouth  and  the  space  just  mentioned  are  widened; 
for  the  pronunciation  of  other  vowels  both  are  contracted;  and  for  others 
one  is  wide,  the  other  contracted.  Admitting  five  degrees  of  size,  both  of 


556  MUSCLE-NERVE    PHYSIOLOGY 

the  opening  of  the  mouth  and  of  the  space  between  the  tongue  and  palate, 
Kempelen  thus  states  the  dimensions  of  these  parts  for  the  following  vowel 
sounds: 

Vowel.          Sound.  Size  of  oral  opening.  Size  of  oral  canah 

a      as  in  "far"  5    3 

a      as  in  "name"  4    2 

e      as  in  "theme"  3    i 

o      as  in  "go"  2    4 

oo      as  in  "cool"  i    5 

Another  important  distinction  in  articulate  sounds  is  that  the  utterance  of 
some  is  only  of  momentary  duration,  taking  place  during  a  sudden  change  in 
the  conformation  of  the  mouth,  and  being  incapable  of  prolongation  by  a  con- 
tinued expiration.  To  this  class  belong  b,  p,  d,  and  the  hard  g.  In  the 
utterance  of  other  consonants  the  sounds  may  be  continuous;  they  may  be 
prolonged,  ad  libitum,  as  long  as  a  particular  disposition  of  the  mouth  and  a 
constant  expiration  are  maintained.  Among  these  consonants  are  h,  m,  n, 
f,  s,  r,  1.  Corresponding  differences  in  respect  to  the  time  that  may  be  oc- 
cupied in  their  utterance  exist  in  the  vowel  sounds,  and  principally  constitute 
the  differences  between  long  and  short  syllables.  Thus  the  a  as  in  far  and 
fate,  the  o  as  in  go  and  fort,  may  be  indefinitely  prolonged;  but  the  same 
vowels  (or  more  properly  different  vowels  expressed  by  the  same  letters), 
as  in  can  and  fact,  in  dog  and  gotten,  cannot  be  prolonged. 

All  sounds  of  the  first  or  explosive  kind  are  insusceptible  of  combination 
with  vocal  tone  (intonation),  and  are  absolutely  mute;  nearly  all  the  con- 
sonants of  the  second  or  continuous  kind  may  be  attended  with  intonation. 

The  tongue,  which  is  usually  credited  with  the  power  of  speech,  plays 
only  a  subordinate,  although  very  important,  part.  This  is  well  shown  by 
cases  in  which  nearly  the  whole  organ  has  been  removed  on  account  of 
disease.  Patients  who  recover  from  this  operation  talk  imperfectly,  and 
their  voices  are  considerably  modified;  but  the  loss  of  speech  is  confined 
to  those  letters  in  the  pronunciation  of  which  the  tongue  is  particularly 
concerned,  namely,  c,  d,  g,  h,  j,  k,  etc. 

Stammering  depends  on  a  want  of  harmony  between  the  action  of  the 
muscles  (chiefly  abdominal)  which  expel  air  through  the  larynx,  and  that  of 
the  muscles  which  guard  the  orifice  (rima  glottidis)  by  which  it  escapes,  and 
of  those  (of  tongue,  palate,  etc.)  which  modulate  the  sound  to  the  form  of 
speech.  Over  either  of  the  groups  of  muscles,  by  itself,  a  stammerer  may 
have  as  much  power  as  other  persons,  but  he  cannot  harmoniously  arrange 
their  conjoint  actions. 


557 


LABORATORY  EXPERIMENTS  ON  MUSCLE  AND  NERVES. 

Physiological  experiments  on  living  muscle  serve  to  demonstrate  many 
of  the  most  fundamental  particulars  of  physiology.  The  muscles  of 
cold-blooded  animals  isolated  from  the  body  retain  their  living  attributes 
for  hours  under  laboratory  conditions.  They  illustrate  practically  all  the 
facts  shown  by  the  muscles  of  warm-blooded  animals. 

i.  The  Use  of  the  Induction  Coil. — The  induced  current  obtained  by 
means  of  an  apparatus  called  an  induction  coil  is  the  most  convenient  and 
reliable  means  of  stimulating  the  muscle  or  the  nerve.  The  arrangement  of 
the  batteries,  keys,  and  coils  for  the  ordinary  muscle  work  is  shown  in  figure 
350.  The  strength  of  the  induced  current  decreases  with  the  distance  of  the 
secondary  coil  from  the  primary,  but  according .  to  a  logarithmic  curve. 
For  ordinary  work  the  centimeter  scale  is  adequate  guide  for  the  proportionate 
strength  of  the  induction  current. 


FIG.  350. — Diagram  illustrating  the  relations  of  the  battery,  keys,  coils,  and  electrodes  as 
used    for    physiological    stimulation. 

At  the  moment  of  closure  of  the  primary  key,  a  current  of  electricity  is 
induced  in  the  secondary  coil,  the  make  current.  The  induced  current  is 
only  momentary  in  duration,  i.e.,  does  not  continue  though  the  primary  circuit 
is  complete.  Also  when  the  primary  current  is  broken  by  opening  the  key  a 
second  current  is  induced,  the  break  current.  The  former  is  in  the  opposite 
direction  to,  the  latter  in  the  same  direction  as  the  primary  current.  In 
ordinary  coils  the  break  current  is  stronger.  The  induction  coil  may  be  used 
to  produce  a  rapid  series  of  shocks  by  means  of  a  magnetic  interrupter,  as  in 
the  Harvard  inductorium. 

a.  Test  the  strength  of  a  series  of  single  make  induction  shocks  beginning 
with  the  weakest  possible  position  of  the  secondary  coil,  and  going  toward  the 
primary  coil  i  cm.  after  each  test.  Test  the  stimulating  effect  by  applying 
the  electrodes  to  the  tip  of  the  tongue.  Record  the  results  as  directed  by  the 
table  below. 


558  MUSCLE-NERVE    PHYSIOLOGY 

b.  Repeat  using  the  break  induction  currents. 


Position  of  the  secondary 
coil  in  centimeters  from 
the  primary 

Relative  stimulating    strength  on  the  tongue,  i.  e.  "none, 
trace,  faint,  mild,  strong,  very  strong,  etc." 

Make  Shock 

Break  Shock 

2.  The  Muscle-nerve  Preparation. — The  classical  muscle-nerve  prep- 
aration is  the  gastrocnemius  muscle  and  the  sciatic  nerve.     Prepare  it 
as  follows :  a.  Kill  the  frog  by  pithing.     This  is  done  by  grasping  the  frog 
firmly  in  one  hand  and  with  the  other  making  a  cut  with  a  blunt  scalpel 
through  the  cranium  just  over  the  medulla,  turning  the  scalpel  so  as  com- 
pletely to  destroy  the  medulla.     Now  take  a  slender  knitting  needle, 
quickly  run  it  up  into  the  cranial  cavity  to  destroy  the  brain,  and  down  the 
spinal  canal  to  destroy  the  cord.     After  a  brief  spasmodic  contraction  of  the 
muscles  of  practically  the  entire  body,  the  frog  remains  limp  and  motion- 
less,    b.  In   making  the  muscle-nerve  preparation  it  is  better  to  isolate 
the  tendon  Achilles  and  insert  its  hook  first,  then  expose  the  nerve,  and 
finally  the  femur.     The  nerve  should  be  prepared  of  as  long  a  length  as 
possible  and  should  not  be  allowed  to  come  in  contact  with  the  skin.     If 
the  preparation  is  to  be  used  in  a  moist  chamber,  the  skin  should  be  entirely 
removed;  if  it  is  to  be  used  in  the  open  air,  the  skin  should  be  left  on.     Use 
care  not  to  stretch  the  nerve,  and  protect  it  from  evaporation.     Cut  the 
femur  long. 

3.  The  Irritability  of  Nerve. — Prepare  a  muscle-nerve  with  its  skin 
on  and  do  not  cut  away  the  foot.     Mount  it  by  inserting  the  femur  in  a 
muscle  clamp,  letting  the  leg  extend  vertically  upward,  and  the  foot  hang 
over.     The  nerve  should  lie  along  the  exposed  moist  femur,  one  end  being 
slightly  free.     Stimulate  the  nerve  in  the  following  ways: 

a.  Electrical  Stimuli. — Apply  the  electrodes  of  the  secondary  coil  of  an 
induction  apparatus  to  the  tip  of  the  nerve.  When  an  induction  current  of 
sufficient  strength  is  produced,  the  muscle  to  which  the  nerve  is  attached 
will  give  contractions,  thus  moving  the  foot.  Notice  that  contractions 
occur  with  both  make  and  break  inductions.  Apply  the  electrodes  from 
the  two  poles  of  a  dry  battery.  When  the  current  of  the  battery  is 
established  a  contraction  will  occur,  but  does  not  continue  during  the  time 
of  the  flow  of  the  current.  When  the  current  is  stopped  a  second  contrac- 
tion occurs.  The  nerve  is  irritable  to  both  galvanic  and  faradic  currents. 


IRRITABILITY    OF    MUSCLE 


559 


b.  Mechanical  Stimuli. — Pinch  the  nerve  lightly  with  forceps,  or  give  it 
a  sudden  stroke  with  the  scalpel  handle.     With  each  mechanical  impact 
there  is  a  single  contraction  of  the  muscle. 

c.  Thermal  Stimuli. — Touch  the  end  of  the  nerve  with  a  glass  rod  heated 
in  boiling  water.     At  each  time  the  nerve  is  brought  in  contact  with  the  rod 
there  will  be  muscular  contraction,  as  in  the  preceding  cases.     The  experi- 
ment succeeds  better  if  the  nerve  comes  in  contact  with  the  rod  for  several 
millimeters  of  length.     If  the  tip  of  the  nerve  has  ceased  to  respond,  then 
snip  it  off  with  the  scissors,  and  repeat  the  experiment  on  the  fresh  end. 

d.  Chemical  Stimulation. — Many  chemical  substances  when  brought  in 
contact  with  living  nerve  fibers  produce  nerve  impulses.     Try  crystals  of 
sodium  chloride,  magnesium  sulphate,  dilute  ammonia,  acetic  acid,  10  per 
cent,  nitric  acid,  i  per  cent,  mercuric  chloride. 

Tabulate  your  observations  on  all  the  forms  of  stimulation  used  above, 
by  the  following  outline: 


Kind  of  stimulation. 

Effect  produced. 

4.  Irritability   of   Muscle. — Repeat    the   experiments   in   number    2 
above,  applying  the  stimuli,  electricity,  etc.,  directly  to  the  muscle  sub- 
stance, choosing  as  far  as  possible  portions  of  muscle  which  do  not  exhibit 
nerve  fiber.     The  muscle  will  usually  respond  by  a  contraction  to  each  of 
the  above  forms  of  stimulation. 

These  tests  do  not  fully  demonstrate  the  direct  irritability  of  muscle 
substance,  since  in  each  case  it  is  possible  that  nerves  may  have  been 
stimulated.  The  nerve  influences  over  the  muscle  can  be  eliminated  by 
the  use  of  drugs,  as  will  be  shown  in  the  next  experiment. 

5.  Independent  Irritability  of  Muscle. — The  influence  01  curara  on 
the  muscle-nerve  preparation  is  demonstrated  as  follows :   Ligate  one  leg  of 
a  frog  near  the  thigh  to  stop  the  circulation  on  that  side.     Now  inject 
under  the  skin  of  the  back  three  drops  of  i  per  cent,  curara,  allowing 
twenty  to  thirty  minutes  for  absorption.     Make  the  following  observations : 
Place  the  animal  on  a  glass  plate  with  back  up  and  dissect  out  the  sciatic 
nerve  in  each  leg.     Use  care  not  to  injure  in  any  way  the  accompanying 
femoral  artery. 

a.  Stimulate  the  muscles  of  the  ligatured  leg,  also  the  muscles  of  the 
curarized  leg,  each  will  contract. 


MUSCLE-NERVE    PHYSIOLOGY 

b.  Stimulate  the  sciatic  nerve  of  the  ligated  leg  above  the  ligature; 
also  the  sciatic  of  the  opposite  side,  both  of  which  have  come  in  contact 
with  the  curara.  Stimulation  of  the  first  nerve  produces  contraction  of  its 
muscle;  of  the  second  nerve  does  not  produce  contraction  of  its  muscle. 

From  this  experiment  of  Claude  Bernard's  it  is  evident  that  the  curara 
does  not  destroy  the  irritability  of  nerve  fiber  nor  the  irritability  of  the 
muscle  fiber,  yet  it  does  destroy  the  influence  of  the  nerve  over  the  muscle, 
probably  acting  as  a  specific  poison  for  the  motor  end-plates.  If  the  motor 
end-plates  are  destroyed,  then  forms  of  stimuli  which  produce  contractions 
of  the  muscle  must  act  directly  on  muscle  substance,  proving  that  muscle 
substance,  as  such,  is  irritable. 


FIG.  351. — Moist  Chamber. 

6.  The  Relation  of  the  Contraction  to  the  Strength  of  the  Stimu- 
lus.— Minimal  and  Maximal  Stimuli. — a.  Prepare  a  recording  cylinder  for 
making  vertical  records  of  the  contractions.  Adjust  the  writing  point  of  the 
muscle  lever  to  the  drum  and  set  the  drum  to  be  moved  by  hand  i  cm.  after 
each  succeeding  contraction.  Set  the  secondary  coil  of  the  induction  ap- 
paratus so  that  it  will  be  too  weak  to  produce  a  stimulus. 

b.  Prepare  a  muscle  nerve  of  the  frog  and  mount  in  the  moist  chamber 
and  arrange  for  stimulating  the  muscle  directly  by  means  of  the  secondary 
current  of  the  induction  coil,  stimulating  apparatus  adjusted  as  in  fig.  350. 

c.  Now  attempt  to  stimulate  the  muscle,  then  move  the  induction  coil 
toward  the  primary  i  cm.  at  a  time  and  repeat  until  the  first  slight  contraction 
appears.     Continue  to  slide  the  secondary  coil  toward  the  primary,  stimulate 
at  each  new  position,  moving  the  drum  forward  for  each  stimulus  as  directed, 
until  a  series  of  contractions  is  obtained  through  the  range  of  variation  of 
induction  of  which  the  apparatus  is  capable,  usually  twenty  or  thirty  con- 
tractions. 


FATIGUE    OF    VOLUNTARY    MUSCULAR    CONTRACTION  561 

A  typical  tracing,  figure  326,  shows  that  as  the  strength  of  the  stimulus  is 
increased  the  amplitude  of  the  contractions  quickly  mounts  from  the  minimal 
to  a  maximal,  after  which  all  further  increase  in  the  strength  of  the  stimulus 
produces  contractions  of  practically  the  same  height.  The  first  perceptible 
contraction  is  called  the  minimal  contraction,  the  strength  of  the  current 
which  produced  it  is  a  minimal  stimulus  for  that  preparation.  The  contrac- 
tions of  the  greatest  amount  are  called  maximal  contractions.  The  weakest 
stimulus  which  produces  a  maximal  contraction  is  called  the  maximal  stimu- 
lus, and  all  stronger  stimuli  supramaximal. 


FIG.  352. — The  Type  of  Contractions  given   by  the   Gastrocnemius  of  the  Frog  to  a 
series  of  stimuli  occurring  at  regular  recurrent  intervals.     (Taskinen.) 

7.  The  Effect  of  Fatigue  on  the  Amplitude  of  a  Series  of  Simple 
Muscle  Contractions. — #,  Arrange  the  recording  apparatus  and  set   the 
induction  coil  for  single  stimuli.     Adjust  the  recording  lever  of  the  muscle 
to  a  smoked-paper  kymograph  and  set  the  speed  of  the  kymograph  to 
revolve  at  the  rate  of  i  mm.  per  second. 

b.  Prepare  a  gastocnemius  muscle  for  direct  stimulation  and  mount  it  in 
a  moist  chamber. 

r.  Now  stimulate  the  muscle  with  the  make  induction  (short-circuiting 
the  break)  once  every  two  seconds.  The  contractions  will  be  recorded  as 
vertical  marks  on  the  drum  in  regular  order,  at  a  distance  of  2  mm.  apart, 
hence  very  slight  changes  in  amplitude  are  readily  detected.  The  contrac- 
tions gradually  increase  in  height  for  the  first  ten  or  twenty  contractions,  the 
phenomenon  of  treppe,  then  run  for  from  fifty  to  one  hundred  contractions 
of  practically  uniform  amplitude,  after  which  there  is  a  gradual  but  sharp 
decrease  known  as  fatigue.  Repeat  on  the  second  gastrocnemius. 

d.  Repeat  the  experiment  after  ten  minutes'  rest.  The  former  varia- 
tions occur  now  very  rapidly,  indicating  that  the  fatigue  effects  are  only 
partially  recovered  from. 

8.  Fatigue   of   Voluntary  Muscular   Contraction,   Demonstration. — 
The  human  voluntary  muscles  are  used  to  demonstrate  this  experiment. 
Use  a  Mosso's  ergograph,  or  any  one  of  its  numerous  modifications.     If  the 
original  form  is  used,  then  the  muscle  should  be  loaded  with  about  3  kilos, 
and  contractions  once  a  second  recorded  until  the  muscle  can  no  longer  lift 
the  load.     The  load  may  have  to  be  adjusted  to  the  individual,  but  should  be 
chosen  so  that  exhaustion  will  be  obtained  with  about  thirty  contractions. 

36 


562  MUSCLE-NERVE    PHYSIOLOGY 

This  experiment  does  not  demonstrate  complete  exhaustion,  but  merely 
fatigue  down  to  a  certain  level.  If  an  apparatus  is  previously  arranged  for 
direct  stimulation  of  the  muscles  by  electric  currents  it  will  be  found  that  the 
contractions  of  the  muscles  still  occur  after  the  voluntary  power  is  lost,  show- 
ing that  at  least  a  part  of  the  phenomenon  of  fatigue,  possibly  the  chief  part, 
is  located  in  the  nervous  tissue  rather  than  in  the  muscle  substance. 

9.  Effect  of  Load  on  the  Height  of  the  Contraction  and  on  the  Work 
of  Voluntary  Muscle. — a.  Make  a  muscle-nerve  preparation  and  arrange 
it  for  stimulation,  as  in  Experiment  6  above.     Set  the  induction  coil  of  the 
stimulating  apparatus  for  an  effective  supramaximal  stimulus.     Record  the 
contractions  as  pairs  of  vertical  lines  on  the  kymograph,  the  pairs  separated 
by  a  distance  of  i  cm.     Begin  with  the  load  of  the  lever  only  for  the  first 
contraction,  then  increase  the  load  by  steps  of  20  grams  each  until  the  muscle 
is  no  longer  able  to  lift  the  weight  used.     Record  two  contractions  for  each 
weight.     Use  care  that  no  mechanical  changes  of  the  apparatus  are  recorded 
so  as  to  confuse  the  record  of  the  contraction  on  the  smoked  cylinder. 

b.  Repeat  the  experiment  on  a  fresh  muscle,  supporting  the  lever  with 
a  tension  of  20  grams  on  the  muscle. 

The  amount  of  work  done  by  the  muscle  at  each  contraction  is  the  prod- 
uct of  the  load  in  grams  times  the  height  in  centimeters.  The  height  of 
the  lift  can  be  obtained  in  this  experiment  from  the  height  of  the  record  on 
the  drum  and  the  lengths  of  the  recording  and  power  arms  of  the  lever,  in 
which  r ./.,  the  length  of  the  recording  lever,  is  to  p.l.,  the  length  of  the  power 
lever,  as  h.,  the  height  of  the  record  obtained,  is  to  c.,  the  actual  shortening  of 
the  muscle.  Compute  the  exact  amount  of  work  done  by  each  contraction 
(c  the  contraction  times  w  the  weight  gives  the  work),  and  tabulate  on  co- 
ordinate paper.  Compare  the  variation  in  work  done  with  the  variation  in 
amplitude  of  the  contraction. 

10.  The  Effect  of  Temperature  on  the  Amplitude  of  the  Muscle  Con- 
tractions.— Prepare  a  muscle-nerve  and  mount  it  in  Porter's  latest  form  of 
temperature  apparatus.     Insert  the  thermometer  bulb  beside  the  muscle. 
Adjust  the  levers  for  vertical  records  on  the  smoked  paper  of  the  kymo- 
graph.    Begin  with  a  temperature  of  tap  water  and  gradually  lower  the 
temperature  of  the  preparation  by  adding  small  amounts  of  crushed  ice  at 
first;  later,  add  ice  and  salt  crystals  to  the  external  chamber,  siphoning  off 
excess  of  fluid  into  a  cup.     Take  care  to  lower  the  external  temperature 
very  slowly  and  gradually,  say  about  one  degree  in  two  minutes.     Stimu- 
late the  muscle  with  a  supramaximal  stimulus  twice  in  rapid  succession, 
for  i°  C.  of  change.     Record  these  contractions  as  pairs  of  vertical  marks 
on   the  drum  i  mm.  apart,  separating  each  pair  by  a  space  of  i   cm. 
When  o°  C.  is  reached,  or  before  if  the  muscle  fails  to  contract  at  a 
higher   temperature,   reverse  the  direction  of  the  temperature  change, 
gradually  but  slowly  increasing  it  until  the  muscle  goes  into  heat  rigor, 
which  begins  at  from  38°  to  40°  C. 


TETANUS  563 

•» 

While  the  muscle  is  entering  rigor,  move  the  drum  i  cm.  for  each  degree, 
as  before,  so  as  to  record  the  development  of  that  process. 

ii.  The  Form  of  the  Simple  Muscle  Contraction. — Striated  muscle 
responds  to  electrical  stimuli  even  though  of  almost  instantaneous  duration. 
The  response  which  the  muscle  gives  to  a  single  stimulus  is  called  a  simple 
muscle  contraction,  and  is  demonstrated  as  follows: 

a.  Arrange  an  induction  coil  with  its  keys,  battery,  and  electrodes  con- 
nected, so  as  to  stimulate  the  muscle  by  the  automatic  key  of  the  Guthrie 


FIG.  353. — Simple  Form  of  Pendulum  Myograph  and  Accessory  Parts.  A,  Pivot  upon 
which  pendulum  swings;  B,  catch  on  lower  end  of  myograph  opening  the  key,  C,  in  its 
swing;  D,  a  spring-catch  which  retains  myograph,  as  indicated  by  dotted  lines,  and  on 
pressing  down  the  handle  of  which  the  pendulum  swings  along  the  arc  to  D  on  the  left  of 
figure,  and  is  caught  by  its  spring. 

apparatus.  Set  the  secondary  coil  at  a  position  which  will  give  a  strong  con- 
traction of  the  muscle,  and  record  this  contraction  on  the  smoked  paper  of  an 
ordinary  recording  cylinder.  Revolve  the  drum  at  a  rapid  rate  using  the 
weight-driven  attachment,  allowing  the  automatic  key  to  be  opened  while  the 
drum  is  turning  at  its  highest  speed.  Or  use  the  pendulum  myograph,  which 
is  especially  constructed  for  this  experiment,  figure  353. 

b.  Make  a  muscle-nerve  preparation  with  the  tendon  isolated  and  the  skin 
removed,  Experiment  2,  and  mount  it  in  a  moist  chamber,  figure  351.  Lay 
the  nerve  across  a  pair  of  platinum  electrodes,  shake  a  little  water  on  the 
inside  of  the  cover  of  the  moist  chamber,  and  place  it  over  the  prepa- 
ration so  as  to  prevent  drying  of  the  nerve  and  of  the  muscle. 


564 


MUSCLE-NERVE    PHYSIOLOGY 


The  muscle  contraction  now  is  recorded  as  a  wave  which  shows  some  con- 
siderable duration  in  time,  figure  354.  Introduce  a  100  double  vibration 
tuning  fork  to  record  the  speed  of  the  drum,  and  take  care  to  mark  the  exact 
point  on  the  record  where  the  automatic  key  is  opened.  (Instead  of  the 
automatic  key  one  may  use  a  hand  key  in  connection  with  a  signal  magnet  to 
mark  the  point  of  stimulation.)  In  this  record  the  muscle  contraction  shows 
three  different  periods  or  phases.  The  first,  a  period  of  no  activity,  called  the 
latent  period,  taking  about  o.oi  of  a  second;  the  second,  the  period  of  rapid 
shortening  known  as  the  contraction  phase,  taking  about  0.04  of  a  second 
on  the  average;  and  the  third,  a  period  of  rapid  relaxation  or  return  to  the 
normal,  which  takes  about  0.05  of  a  second,  see  figure  354. 

The  time  and  character  of  the  simple  muscle  contraction  will  be  influ- 
enced by:  a,  load  or  tension;  b,  the  exact  temperature;  c,  by  the  amount  of 
work  it  has  previously  done,  or  fatigue ;  d,  by  the  time  since  it  was  isolated 
from  the  circulation.  Perform  a  series  of  experiments  varying  these 
effects,  and  record  the  results  by  the  following  outline: 


Number  of 
the 
Experiment. 

Muscle 
Used. 

Temper- 
ature. 

Load. 

Total  Time  of 
Contraction, 
in  Seconds. 

Latent 
Period,  in 
Seconds. 

Contraction 
Period,  in 
Seconds. 

Relaxation 
Period,  in 
Seconds. 

FIG.  354. — Record  of  a  Simple  Contraction  of  the  Gastrocnemius  of  the  Frog.  Time 
in  .01  of  a  second.  St,  Moment  of  stimulation.  Record  taken  on  a  rapid  drum  that  was 
provided  with  an  automatic  key. 

12.  The  Effect  of  Fatigue  on  the  Time  of  the  Simple  Contraction. 
Prepare  a  muscle  nerve  and  mount  it  in  the  moist  chamber,  arrange  for  the 
record  as  directed  under  5  above.  Make  a  series  of  records  of  the  simple 
contraction  when  automatically  stimulated,  recording  only  every  tenth  or 
twentieth  contraction — the  intermediate  contractions  should  be  shunted  and 
are  used  merely  to  produce  fatigue.  After  a  time  the  contractions  will  not 
only  diminish  in  amplitude,  but  there  will  be  a  gradual  increase  in  the  time 


TETANUS  565 

consumed  by  the  contraction.  This  increase  in  time  falls  very  slightly  on 
the  latent  period,  is  more  pronounced  in  the  contraction  phase,  but  is  very 
marked  in  the  relaxation  phase,  figure  355. 


FIG.  355. — Contractions  of  the  Gastrocnemius  Muscle  to  Show  Fatigue.     The  numbers 
printed  on  the  figure  indicate  the  contraction  in  the  series  which  is  recorded.     (Lee.) 

13.  The  Effect  of  Temperature  on  the  Time  of  the  Simple  Con- 
traction.— Repeat  Experiment  10  but  record  the  contractions  by  the  method 
described  in  Experiment  1 1  above,  recording  a  contraction  for  every  change 
of  5°  C.     Measure  the  time  and  amplitude  of  the  different  contractions, 
and  the  phases  of  the  simple  contractions,  and  tabulate  them  as  shown  in 
Experiment  n. 

14.  Tetanus. — A   continued   contraction   of   a   voluntary   muscle   can 
be  shown  to  be  a  fusion  of  simple  muscle  contractions.     This  is  called  a  teta- 
nus.    Arrange  the  induction  coil  for  stimulating  with  a  series  of  rapidly 
repeated  stimuli.     Set  the  secondary  coil  so  the  break  inductions  only  will 
stimulate  thus  securing  one  stimulus  per  vibration  of  the  tetanometer  key. 
The  rate  of  the  stimulation  is  obtained  from  the  Harvard  tetanometer,  a  form 
of  key  for  rapidly  interrupting  the  current,  which  should  be  connected  with 
the  primary  coil  instead  of  the  key,  K,  figure  350.     Prepare  a  muscle-nerve 
in  the  moist  chamber  and  stimulate  the  muscle  at  a  rate  of  10  per  second, 
record  the  contractions  on  the  drum  moving  at  a  speed  of  about  2  cm.  per 
second.     Use  care  not  to  overfatigue  the  muscle,  i.e.,  stimulate  it  only  2  sec- 
onds at  a  time  controlling  the  time  by  the  extra  key  in  the  primary  circuit. 
Repeat  this  test,  increasing  the  rate  of  stimulation  each  time  by  5,  that  is 
stimulate  at  10,  15,  20,  etc.,  per  second.    In  the  first  stimulation  there  will  be 
a  series  of  simple  contractions  with  almost  complete  intervening  relaxations. 
As  the  rate  is  increased  these  relaxations  become  less  and  less  until  presently 
a  rate  is  found  which  produces  continuous,  apparently  uninterrupted  contrac- 
tions.    This  is  a  tetanus.     The  others  are  incomplete  tetani.     The  frog's 
gastrocnemius  at  a  temperature  of  20°  C.  is  tetanized  with  a  stimulation 
of  from  25  to  30  per  second. 


566 


MUSCLE-NERVE    PHYSIOLOGY 


15.  DuBois-Reymond's  Law  of  Galvanic  Stimulation. — Connect  a  single 
dry  cell,  a  key  switch,  and  a  meter-rheocord  in  circuit.     To  the  zero  end 
of  the  rheocord  attach  a  lead-off  through  a  pole-changer  to  one  of  the 
non-polarizable  boot  electrodes  of  a  Harvard  moist  chamber.     Attach 
the  free  wire  of  the  lead-off  to  the  other  boot  electrode.     Make  a  muscle- 
nerve  preparation  with  a  long  nerve.     Mount  in  a  moist  chamber  so  that 
the  nerve  will  lie  across  the  two  boot  electrodes,  one  quite  close  to  the 
muscle  but  not  allowed  to  touch  it,  and  the  other  as  far  from  the  muscle 
as  the  length  of  the  nerve  will  permit.     Record  on  a  drum  but  move  the 
drum  by  hand. 

The  galvanic  current  (when  of  moderate  intensity)  stimulates  only  at 
the  time  of  changing  intensity,  i.e.,  on  making  and  on  breaking  the  current. 
Connect  the  free  wire  of  the  lead-off  with  the  rheocord  between  the  70 
and  zoo  cm.  points.  The  exact  point  will  vary,  the  stronger  the  battery 
the  nearer  to  point  of  attachment  to  the  70  cm.  mark.  Note  the  responses 
of  the  muscle,  (a)  on  closing  the  circuit,  (b)  during  the  flow  of  the  current, 
and  (c)  at  the  breaking  of  the  circuit.  Make  records  covering  these  facts 
known  as  DuBois-Reymond's  Law  of  Stimulation. 

1 6.  Pfluger's  Law  of  Irritability. — Both  strength  of  current  and  direc- 
tion of  flow  of  current  are  factors  in  the  determination  of  the  physiological 
effects  of  the  galvanic  current  on  irritability.     This  law  is  tabulated  as 
follows : 


Strength  of  galvanic 
current 

Ascending    current 

Descending   current 

Making 

Breaking 

Making 

Breaking 

Very  weak  currents  
Medium  currents  
Very  strong  currents  

Contraction 
Contraction 
No  contraction 

No  contraction 
Contraction 
Contraction 

Contraction 
Contraction 
Contraction 

No  contraction 
Contraction 
No  contraction 

Determine  the  direction  of  flow  of  galvanic  current  in  your  apparatus 
and  demonstrate  the  several  parts  of  Pfluger's  Law  as  given  in  the  table. 
Begin  with  the  very  weak  ascending  currents,  use  one  battery  with  the 
lead-off  attached  between  20  and  40  cm.  for  very  weak  currents.  Reverse 
the  direction  of  the  current  by  the  pole-changer.  Use  two  batteries  with 
lead-off  between  50  and  80  cm.  for  medium  currents;  and  two  batteries 
with  lead-off  at  90  cm.  for  very  strong  current,  or  four  batteries  with  lead- 
off  at  80  cm.  Make  records  of  each  contraction  and  label  as  to  its  type 
and  direction. 

17.  Electrotonus. — The  change  in  irritability  at  the  physiological  poles 
of  a  nerve  through  which  a  galvanic  current  is  passing  is  called  electro- 
tonus.  Mount  a  fresh  muscle  nerve  preparation  in  a  Harvard  moist 


THE    ACTION    CURRENT    OF    MUSCLE  567 

chamber  with  a  long  nerve  and  nonpolarizable  electrodes.  Use  a  small 
platinum  stimulating  electrode  arranged  to  stimulate  the  nerve  at  differ- 
ent points,  a,  on  the  muscle  side  of  the  boot  electrode,  b,  next  to  the  muscle 
but  as  close  to  the  boot  electrode  as  possible.  Use  single  induction  cur- 
rents for  stimulation  of  the  nerve  during  the  flow  of  a  galvanic  current  as 
in  Experiment  17. 

Record  and  check  the  change  in  nerve  irritability  during  electrotonus 
against  the  diagram,  figure  335; 

1 8.  The  Action  Current  of  Muscle,    a.  The  Rheoscopic  Frog. — The 
action  current  produced  by  a  single  contraction  of  a  gastrocnemius  muscle 
is  enough  to  stimulate  the  sciatic  nerve  of  a  second  preparation  when 
the  second  nerve  is  laid  across  the  first  muscle  in  such  a  way  as  to  be  a 
conductor  of  its  action  current.     Expose  the  two  gastrocnemius  muscles 
of  a  frog  pinned  horizontally  to  a  frog  board.     These  need  not  be  separated 
from  the  skeleton.     Bring  the  two  near  to  each  other  and  pin  the  knees 
to  the  frog  board.     Attach  the  tendon  of  the  second  to  a  lever  and  bring 
its  long  sciatic  nerve  in  contact  with  the  tendon  and  the  belly  of  the  first 
muscle.     Now  stimulate  the  nerve  of  the  first  muscle.     When  the  first 
muscle  contracts  in  response  to  the  nerve  impulse  from  its  nerve,  the 
action  current  developed  will  flow  through  the  loop  of  the  second  nerve  in 
physical  contact  only  with  the  first  muscle.     Stimulation  of  the  nerve  is 
indicated  by  the  contraction  of  the  second  muscle  as  recorded  on  the 
drum. 

b.  By  careful  manipulation  the  "current  of  injury"  or  "demarcation 
current"  which  develops  in  the  first  muscle  when  cut  across  the  middle 
will  also  stimulate  the  second  nerve.     Use  a  glass  rod  to  adjust  the  nerve. 
Bring  the  middle  of  the  nerve  in  contact  with  the  proximal  tendon  of  the 
cut  muscle,  then  suddenly  drop  the  free  end  of  the  nerve  onto  the  cut  end  of 
the  muscle.     The  nerve  will  be  stimulated  and  the  second  muscle  will 
contract. 

c.  Action  Current  of  the  Heart. — Prepare  a  perfectly  fresh  gastrocnemius 
with  a  long  sciatic.     Expose  a  large  turtle's  heart.     Support  the  sciatic 
by  a  glass  rod  so  that  it  touches  the  heart  at  the  base  and  the  apex  only. 
The  cardiac  action  current  will  stimulate  the  sciatic  at  each  contraction. 

19.  Unipolar  Stimulation. — The  stimulating  effect  of  an  electric  cur- 
rent is  proportional  to  the  intensity  of  the  current  per  unit  area  of  the 
physiological  electrode.     When  one  electrode  is  made  indifferent  by  spread- 
ing the  current  over  a  large  surface,  then  the  stimulations  will  be  limited 
to  the  smaller  electrode.     Use  the  Harvard  apparatus,  bind  the  indifferent 
electrode  on  the  neck  and  with  the  other  verify  the  motor  points  in  the 
arm  shown  in  figure  336. 

This  method  can  be  used  to  demonstrate  DuBois-Reymond's  law  on 
human  nerves.  Apply  the  stimulating  electrode  over  the  median  nerve 
at  the  elbow.  Stimulation  with  a  medium  current  will  occur  only  on  clos- 


568 


MUSCLE-NERVE    PHYSIOLOGY 


ing  the  descending  current  or  on  opening  the  ascending  current.  On 
reversing  the  current  the  opening  stimulus  will  not  be  effective. 

20.  Cardiac  Muscle. — Cardiac  muscle  differs  from  voluntary  in  that 
the  contractions  occur  rhythmically  and  automatically.  This  is  shown  by 
the  isolated  frog's  heart,  which  continues  to  contract  when  bathed  with 
blood  or  salt  solution,  often  for  hours.  This  isolated  heart,  however,  has  a 
complicated  local  nervous  mechanism.  The  apex  of  the  ventricle  of  the 
terrapin's  heart  is  practically  free  from  nerve  ganglia  and  is  used  to  demon- 
strate the  characteristics  of  pure  cardiac  muscle. 

a.  Cut  a  strip  off  the  apex  of  the  terrapin's  ventricle,  as  shown  in  figure 
214,  and  mount  it  by  means  of  light  silk-thread  ligatures  tied  around  the  two 
ends  of  a  strip  and  attached  to  the  apparatus  shown  in  figure  356.  The  loop 
of  the  lower  thread  should  be  i  cm.  long,  of  the  upper  thread  long  enough  to 


FIG.  356. — Arrangement  of  Apparatus  for  Studying  the  Contractions  of  the  Strip  of  the 

Apex  of  the  Ventricle. 

reach  to  the  lever  above.  When  such  ventricular  strips  are  immersed  in 
ordinary  0.7  per  cent,  sodium  chloride  they  will  begin  contractions  in  a  few 
minutes,  twenty  minutes  or  so.  The  contractions  are  regular  in  rate  and  will 
continue  through  two  or  three  hours,  gradually  becoming  smaller  and  smaller 
until  a  standstill  is  reached.  If  the  strip  is  immersed  in  its  own  serum  it  will 
give  only  occasional  contractions,  but  it  remains  irritable  and  capable  of 
contracting  at  any  moment.  If  changed  to  salt  solution,  the  salt  solution 
apparently  brings  out  the  automatic  rhythm  by  an  increase  in  its  irritability. 
b.  Portions  of  the  auricle  and  of  the  sinus,  especially  the  latter,  are  more 
highly  rythhmic  than  portions  of  the  ventricle,  due  to  a  specific  difference  in 
the  muscle  cells  themselves  rather  than  to  the  nervous  mechanism  contained. 
Prepare  a  strip  from  the  auricle  and  compare  it  with  the  ventricle. 


CARDIAC    MUSCLE  569 

c.  Repeat  with  sinus  strips  using  care  to  compensate  the  weight  of  the 
lever. 

Refer  to  the  experiments  on  cardiac  muscle  at  the  end  of  the  chapter  on 
Circulation. 

21.  Involuntary  Muscle,  Coldblooded  Animal. — Strips  of  smooth  or 
involuntary  muscle  cut  from  the  stomach  of  the  frog  will  show  the  physio- 
logical reactions  of  this  type  of  muscle.  Mount  a  spiral  strip  of  frog 
stomach  muscle  in  the  Harvard  warm  chamber  holder.  Use  a  wire  to 
connect  the  muscle  with  the  lever  as  in  figure  356.  Take  care  not  to 
load  too  heavily,  the  ordinary  muscle  lever  is  often  too  great  a  tension. 
Mount  a  signal  magnet  to  automatically  mark  the  time  of  stimulation. 

a.  Frog  stomach  muscle  will  develop  automatic  contractions  in  a  few 
moments.     Record  through  two  or  three  circuits  of  the  drum.     The  speed 
should  be  i  mm.  per  second. 

b.  If  automatic  contractions  do  not  develop  stimulate  the  muscle 
by  interrupted  currents  of  varying  strength.     After  a  variable  latent 
period  contractions  will  develop.     If  the  automatic  contractions  are  occur- 
ring, stimulation  will  increase  the  amount  and  sometimes  the  frequency 
of  the  contractions. 


FIG.  3560. — Figure  Showing  the  Type  of  Contraction  of  a  Strip  of  Muscle  from  the 
Stomach  of  a  Frog.  The  muscle  was  stimulated  with  an  interrupted  current  during  the 
time  indicated  by  the  signal  tracing,  immediately  below  the  time  tracing.  Time  in  seconds. 

c.  A  strip  of  terrapin  stomach  muscle   will  not  develop  automatic 
rhythm  but  will  respond  to  comparatively  strong  stimulation.     Increasing 
the  temperature  to  30°  will  often  initiate  rhythm. 

d.  Smooth  muscle,  like  voluntary  muscle,  responds  to  variation  in 
temperature,  fatigue,  strength  of  stimulus,  etc.     It  is  more  sensitive  to 
injury.     It  shows  a  contraction  amplitude  proportional  to  the  stimulus. 


570  MUSCLE-NERVE    PHYSIOLOGY 

It  is  kept  in  better  condition  in  an  atmosphere  of  oxygen.  Test  as  many 
of  these  points  as  time  permits. 

e.  Moisten  the  strip  with  i  per  cent,  barium  chloride. 

/.  Mount  a  section  of  the  oviduct  of  the  turtle.  This  tube  displays 
a  remarkably  even  and  uniform  but  slow  automatic  rhythm. 

22.  Periodic  Contractions  of  the  Rat's  Uterus. — A  splendid  preparation 
of  smooth  muscle  in  a  warmblooded  animal  is  obtained  by  splitting  the 
rat's  uterus  longitudinally,  giving  one  fallopian  tube  and  one-half  of  the 
uterus  as  a  preparation. 

a.  Mount  a  rat's  uterus-fallopian  tube  in  the  Harvard  warm  chamber 
and  immerse  in  a  bath  of  oxygenated  Ringer's  solution  at  37°  C.     Rhyth- 
mic   contractions    of    a    comparatively    uniform    amplitude    will    begin 
promptly  and  continue  through  a  long  interval. 

b.  While  the  preparation  is  contracting  rhythmically  add  to  the  20 
cc.  of  Ringer's  solution  i  cc.  of  .01  per  cent,  epinephrin.     This  hormone 
stimulates  the  autonomic  nerve  endings  producing  inhibition.     The  effect 
passes  away  after  a  few  minutes  or  on  renewing  the  solution. 

A  similar  preparation  can  be  made  from  the  uterus  of  a  guineapig, 
cat  or  rabbit.  Epinephrin  stimulates  the  inhibitory  mechanism  of  the 
virgin  uterus,  but  produces  contraction  of  the  muscle  of  the  gravid  uterus. 

23.  Ciliary  Contractions. — Ciliated  Epithelium. — Make  a  preparation 
of  ciliated  epithelium  by  cutting  out  the  esophagus  of  a  terrapin  or  frog, 
slitting  it  open  longitudinally,  and  smoothing  it  out  on  a  cork  block.     The 
cilia  of  this  membrane  will  drive  in  the  direction  down  the  esophagus.     Test 
the  rate  at  which  different  loads  are  moved  and  measure  the  distance  on  the 
preparation  as  follows:     Cut  pieces  of  clean  white  paper  about  4  and  6  mm. 
square.     Select  a  favorable  area  on  the  ciliated  surface  as  long  as  possible, 
place  the  4  mm.  square  paper  at  the  beginning  of  the  area,  and  measure  the 
time  which  it  takes  to  travel  the  distance.     Measure  the  speed  in  terms  of 
seconds  per  centimeter.     Now  replace  the  paper  at  the  point  of  beginning 
and  load  it  with  small  weighed  cubes  of  paraffin.     The  rate  at  which  the 
load  is  carried  will  slightly  increase  at  first  as  the  load  is  increased,  but  later 
will  sharply  decrease.     Elevate  one  end  of  the  ciliated  membrane  and  repeat 
the  experiment  with  different  loads  so  that  the  cilia  will  now  carry  the  load 
uphill.     Calculate  the  work  done  in  terms  of  gramcentimeters  of  work  per 
square  centimeter  of  ciliated  surface  acting  on  the  load. 

24.  Rate  of  the  Nerve  Impulse. — Prepare  a  muscle-nerve  of  a  large 
frog  with  the  entire  sciatic  nerve.     Sever  the  nerve  where  it  leaves  the  cord, 
leaving  it  attached  to  the  fascia.     Mount  in  a  moist  chamber  using  two  pairs 
of  electrodes.     Set  one  pair  of  electrodes  on  the  nerve  as  near  the  muscle  as 
possible,  the  other  at  the  extreme  end  of  the  sciatic.     Take  several  pairs  of 
simple  muscle  contractions  on  the  pendulum  myograph  stimulating  in  order, 
i,  the  long  nerve,  and  2,  the  short  nerve.     The  latent  periods  must  be  marked 


RATE  OF  THE  NERVE  IMPULSE  571 

with  greatest  accuracy  and  made  as  long  as  possible.  The  difference  in  the 
time  of  the  two  latent  periods  of  each  pair  represents  the  time  required  for  the 
nerve  length  between  the  two  pairs  of  electrodes.  Compute  the  rate  of  the 
nerve  impulse  in  meters  per  second.  Make  a  table  of  all  the  tests  recorded 
and  draw  averages. 

25.  Relation  of  Total  Work  to  Nutrition  of  Muscle. — Pith  a  frog  and 
pin  it  prone  on  a  frog  board.     Isolate  one  gastrocnemius  and  measure  its 
total  work  as  in  experiment  7.     Inject  2  cc.  of  one  per  cent,  glucose  in 
Ringer  into  the  circulation,  or  perfuse  the  heart  directly  with  Ringer's 
solution  containing  0.2  per  cent,  glucose.     Now  repeat  the  measurement 
of  total  work  on  the  second  gastrocnemius  muscle.     The  experiment  will 
be  more  striking  if  the  records  are  made  parallel  on  the  same  paper  with  the 
drum  at  constant  speed. 

Variations  of  this  demonstration  can  be  made  by  comparing  the  work 
of  the  two  muscles  of  a  frog,  one  while  the  normal  circulation  continues, 
the  other  with  the  circulation  tied  off  by  ligature  around  the  thigh. 

26.  Production  of  Carbondioxide  During  Contraction. — Prepare  a 
CO2-absorbing  apparatus.     Charge  it  with  Ringer's  solution  containing 
phenolphthalein   indicator.     Reduce  the  hydrogen  iron   content   to   a 
constant,  as  indicated  by  the  color  of  the  solution  matched  against  a 
pink  standard,  by  aerating  with  CC>2  free  air.     Prepare  a  small  muscle 
and  attach  it  to  the  holding  device.     Insert  in  the  apparatus,  and  accu- 
rately measure  the  time  until  the  color  of  the  indicator  is  just  discharged. 
This  measures  the  rate  of  CO2  production  in  the  resting  muscle.     Repeat 
until  the  readings  are  constant. 

Now  make  the  test  while  stimulating  the  muscle  to  a  mild  tetanus. 
The  apparatus  and  solution  are  to  be  standardized. 


CHAPTER  XTV. 
THE  NERVOUS  SYSTEM. 

The  nervous  system  consists  of  an  extremely  complex  anatomical  mass 
of  nerve  cells  and  fibers.  It  is  usually  described  as  made  up  of  two  main 
divisions,  the  cerebro-spinal  system  and  the  sympathetic.  These  two  divi- 
sions must  be  regarded  as  parts  of  one  great  whole,  and  in  no  sense  coordinate. 
The  gross  subdivision  of  the  nervous  system  may  be  given  as  the  following: 
First,  the  cerebro-spinal  axis,  which  consists  of  matter  within  the  bony  cra- 
nium and  spinal  column,  constituting  the  brain  and  cord.  Second,  smaller 
masses  for  the  most  part  in  the  abdominal  and  thoracic  cavities,  also  in  the 
neck  and  head,  and  constituting  the  sympathetic  or  autonomic  ganglia. 
Third,  the  nerves  or  bundles  of  nerve  fibers  which  connect  the  central 
nerve  axis  with  the  periphery  and  with  -the  sympathetic  ganglia.  Fourth, 
there  are  special  peripheral  organs  in  connection  with  the  beginnings  and 
endings  of  the  nerve  fibers,  the  one  for  receiving  nerve  stimuli  the  receptors, 
the  other  for  transmitting  stimuli  to  other  tissues,  the  effectors.  These 
are  properly  parts  of  the  nervous  system.  The  peripheral  organs  for 
receiving  stimuli  constitute  the  sense  organs  and  will  be  treated  in  a  sepa- 
rate chapter. 

FUNCTION  OF  THE  NERVE  CELL. 

The  Nerve  Cell. — The  nerve  cell,  the  neurone,  is  considered  the  ana- 
tomical and  physiological  unit  of  the  nervous  system.  Waldeyer  introduced 
the  term  neurone  to  designate  the  nerve  cell  body  and  all  its  constituent  proc- 
esses. Much  of  the  details  of  structure  of  types  of  neurones,  both  as  to  the 
structure  of  the  cell  body  and  all  its  processes,  have  already  been  given  in 
Chapter  II.,  see  figures  83  to  106.  It  is  sufficient  to  recall  that  the  types  of 
nerve  cells  found  in  various  parts  of  the  nervous  system  vary  extremely. 
The  peculiar  feature,  however,  consists  in  the  fact  that  the  cell  body  has 
one  or  more  processes,  branches,  or  arborizations,  see  figure  365.  Some- 
times these  processes  are  short  but  complexly  branched,  sometimes  they  are 
exceedingly  long  as  compared  with  the  extent  of  the  cell  body.  The  cell 
processes  may  or  may  not  be  medullated.  In  medullated  nerves  the  ad- 
ventitious structures  are  subdivided  into  nodes,  but  the  axis-cylinder  process 
is  to  be  regarded  as  a  continuation  of  the  protoplasm  of  the  cell  body. 

In  recent  years  the  structure  of  the  cell  body  and  its  branches  has  been 
very  carefully  investigated,  with  the  result  that  we  have  learned  that  the 
intimate  structure  is  very  complex.  Networks  of  neurofibrillae  have  been 

572 


THE    NEURONE    THEORY 


573 


described  not  only  in  the  cell  body,  but  extending  throughout  the  course  of 
the  processes  and,  in  fact,  from  cell  to  cell.  We  are  not  in  a  position  at  the 
present  time  fully  to  determine  what  bearing  these  neurofibrillae  have  on  our 
accepted  theories  of  nerve  function,  other  than  that  they  are  assumed  to  be 
the  conducting  elements. 


13 


FIG.  357. — Morphological  changes  of  nerve  cells  during  function  activity  and  during 
recuperation,  i.  Normal  resting  Purkinje  cell.  2  to  9.  The  main  stages  in  progressive 
activity.  10  to  13.  The  regular  course  of  recovery.  (Dolley.) 


The  Neurone  Theory. — Our  knowledge  of  the  function  of  the  ner- 
vous system  is  best  explained  on  the  basis  of  the  neurone  theory,  which  con- 
siders the  neurone  as  a  physiological  unit.  By  this  view  each  gross  divi- 
sion of  the  nervous  system  is  supposed  to  consist  of  a  large  number  of  in- 


574 


THE    NERVOUS    SYSTEM 


dividual  neurones,*  each  of  which  is  a  more  or  less  complete  morphological 
unit  capable  of  Carrying  on  certain  functions  of  its  own.  Each  of  these 
neurones  maintains  physiological  continuity  with  its  associates,  presumably 


lisi 


B 


^FiG.  358. — Spinal  Ganglion  Cells  from  the  Cat.  A,  Normal  taken  before  stim- 
ulation; B,  taken  after  five  hours'  stimulation.  From  the  right  and  left,  eight  thoracic 
ganglia.  (Hodge.) 

by  protoplasmic  contact  rather  than  by  continuity;  so  that  well-marked  paths 
of  conduction  are  possible  throughout  the  extent  of  the  particular  mass  of 

*  "According  to  the  estimations  of  Meynert,  the  cortex  of  the  cerebral  hemisphere 
alone  contains  twelve  hundred  millions  of  ganglion  cells.  Donaldson  (The  Growth  of  the 
Brain,  a  Study  of  the  Nervous  System  in  Relation  to  Education,  i2mo,  London,  1897, 
P-  J59)  states  that  for  the  total  number  of  nerve  cells  in  the  central  nervous  system  three 
thousand  millions  is  a  moderate  estimate."  (Barker,  the  Nervous  System  and  its  Con- 
stituent Neurones,  p.  42,  1899.) 


WALLERIAN   DEGENERATION  575 

which  the  neurone  is  a  part,  and  throughout  the  adjacent  masses.  By  this 
view,  paths  of  conduction  are  made  up  of  series  or  chains  of  individual  neu- 
rones which  are  in  physiological  continuity. 

The  Characteristics  of  the  Individual  Neurone. — The  function  of 
the  nerve  cell  may  be  discussed  under  two  headings:  The  function  of  the 
cell  body,  and  the  function  of  the  cell  processes. 

The  cell  body  of  the  neurone  is  the  part  that  contains  the  nucleus  and  is 
the  center  of  those  activities  which  influence  the  metabolism  of  the  cell  itself. 
If  the  cell  body  be  isolated  from  its  processes,  the  processes  will  degenerate, 
while  the  body  continues  to  live.  In  other  words,  the  cell  body  may  be  con- 
sidered as  the  center  of  those  trophic  influences  which  regulate  the  nutrition 
of  the  processes.  Although  the  nerve  cell  as  a  whole  is  in  many,  perhaps  in 
most,  cases  a  conducting  organ,  still  those  physiological  processes  which  go  on 
in  it  produce  marked  changes  in  the  protoplasm  of  the  cell  body.  Hodge 
has  demonstrated  that  nerve  cells  which  have  been  active  for  several  hours, 
in  case  of  sparrows  which  have  been  flying  about  actively  throughout  the  day, 
or  in  bees  after  a  day's  work,  show  marked  evidences  of  fatigue.  These 
evidences  consist  in  the  decrease  in  the  size  of  the  nucleus  and  the  appearance 
of  vacuoles  in  its  structure,  also  in  the  shrinking  of  the  protoplasm  of  the  cell, 
which,  in  case  of  the  cells  of  the  spinal  ganglia,  draws  away  from  its  capsule. 
If  the  cells  are  examined  early  in  the  morning  the  fatigue  changes  will  not 
be  present,  the  cell  having  recuperated  during  the  period  of  rest  at  night.  It 
has  also  been  found  that  the  Nissl  granules  which  are  present  in  the  cell  body 
of  resting  cells  decrease  in  number  and  show  evidence  of  disintegration  in 
cells  that  have  been  stimulated  for  several  hours,  or  that  have  been  in  pro- 
longed functional  activity.  Figure  357  presents  the  changes  in  nerve  cell 
structure  which  can  be  determined  from  microscopical  preparations  after 
various  stages  of  functional  activity.  In  this  series  worked  out  by  Dolley  on 
the  dog,  i  to  9  show  these  changes.  Progressive  recuperation  is  shown  in 
figures  10  to  13.  As  a  matter  of  fact,  the  different  individual  brain  cells 
are  subjected  to  different  degrees  of  activity  during  prolonged  physiological 
strain.  Hence  all  these  cells  are  found  in  one  and  the  same  preparation,  the 
relative  number  varying  however. 

Hodge  studied  the  changes  in  spinal  ganglion  cells  induced  by  artificial 
stimulation.  The  results  are  shown  pictorially  in  figure  358. 

The  nerve  processes  or  fibers  are  primarily  conducting  structures.  But 
fibers  are  susceptible  to  artificial  stimulation,  as  shown  in  the  previous 
chapter;  that  is,  they  are  irritable.  They  are  influenced  by  certain  changes  in 
the  environment,  but  they  do  not  show  evidence  of  fatigue  upon  prolonged 
functional  activity. 

Nutritive  Influence  of  the  Cell  Body  over  its  Processes — Wallerian 
Degeneration. — The  control  of  the  cell  body  over  the  nutrition  of  the 
cell  processes  is  demonstrated  by  the  changes  which  occur  when  these  proc- 


576  THE    NERVOUS    SYSTEM 

esses  are  severed  from  connection  with  the  cell  body.  Under  such  conditions 
the  axis-cylinder  process  completely  degenerates.  Howell  and  Huber  have 
followed  the  degenerative  changes  in  medullated  nerve  fibers.  The  medul- 
lated  fiber  in  the  course  of  three  or  four  days,  in  mammals,  breaks  up  into 
elliptical  segments  of  myelin,  containing  small  fragments  of  the  axis-cylinder. 
These  changes  in  the  cut-off  section  of  nerve  occur  simultaneously  through- 
out its  whole  extent.  In  the  course  of  a  few  weeks  regenerative  changes  be- 
gin, apparently  under  trophic  influence  of  the  nuclei  of  the  primitive  sheath. 
These  nuclei  increase  in  number  and  form  small  masses  of  protoplasm  which 
ultimately  produce  a  strand  of  embryonic  protoplasm,  which  is  described  as 
the  "band  fiber."  If  the  ends  of  the  sectioned  nerve  have  originally  been 
brought  together  and  sutured  in  place,  then  the  axis-cylinder  processes  of  the 
portion  of  the  nerve  fiber  still  attached  to  the  cell  body  will  grow  down  into 
the  peripheral  fibers,  thus  forming  new  axis-cylinder  processes  along  the 
course  of  the  band  fiber.  If  the  stumps  of  the  nerves  are  not  so  brought  to- 
gether, then  apparently  the  band  fiber  again  degenerates,  especially  in  adult 
tissues,  though  it  has  been  claimed  by  Bethe  and  others  that  complete  regen- 
eration of  the  peripheral  fiber  will  take  place  in  very  young  animals.  Even  if 
complete  regeneration  should  take  place  in  the  peripheral  fiber,  unless  con- 
nection were  established  between  it  and  the  central  end  of  the  fiber  it  would 
ultimately  disintegrate  and  could  only  temporarily  carry  on  any  physio- 
logical function.  Complete  regeneration  requires  from  three  months  to 
two  years. 

The  central  end  of  the  divided  nerve,  that  is,  the  part  maintaining  con- 
nection with  the  cell  body,  usually  degenerates  for  a  few  nodes  only,  then  re- 
generation and  growth  of  the  original  stump  proceed.  Instances  are  observed 
in  certain  cases  where  the  degeneration  of  the  entire  central  fiber,  includ- 
ing its  cell  body,  takes  place.  This  happens  particularly  in  those  relations 
where  the  original  neurone  forms  a  link  in  a  conducting  path.  Such  a  con- 
ducting neurone  would  no  longer  perform  its  proper  function  so  would 
atrophy  just  as  would  a  muscle  fiber  when  cut  off  from  its  nervous  relations. 

In  conclusion,  one  may  infer  that  the  cell  body  exercises  a  nutritive  or 
trophic  control  over  the  protoplasm  of  its  branches,  just  as  we  have  already 
seen  the  neurone  as  a  whole  exercises  trophic  control  over  the  nutritive 
processes  taking  place  in  the  tissue  to  which  its  branches  are  distributed,  for 
example,  the  voluntary  muscles. 

Specific  Energy  of  the  Nerve  Impulses. — We  have  already  discussed 
the  fact  that  a  nerve  fiber,  also  its  cell  body,  is  irritable  to  various  forms  of 
mechanical,  electrical,  and  other  stimuli.  In  the  complex  of  activity  of  the 
nervous  system  it  is  found  that  whatever  the  form  of  the  external  stimulus 
applied  to  a  nerve,  the  resulting  nerve  impulse  produces  the  same  effects 
in  the  central  nervous  system.  The  reaction  in  consciousness  is  constant 
and  unvarying  and  independent  of  the  character  of  the  external  stimulus. 


TRANSMISSION    OF    NERVE    IMPULSES    THROUGH   THE    NEURONE     577 

For  example,  if  the  temperature  points  on  the  skin  be  stimulated,  as  they 
may,  by  a  number  of  widely  different  types  of  external  stimuli,  the  result- 
ing sensation  is  the  same;  stimulation  of  a  "heat  point"  by  ice  produces  a 
sensation  of  heat,  not  cold.  This  phenomenon  has  been  called  the  specific 
energy  of  the  nerve  impulse,  and  the  term  was  first  advanced  by  Johannes 
Miiller.  Different  views  are  presented  in  explanation.  But  it  seems  ra- 
tional to  believe  that  the  gist  of  the  matter  rests  in  two  factors:  i.  The 
highly  differentiated  sense  organ  is  adapted  especially  to  stimulation  by  a 
particular  stimulus,  as  the  eye  by  light.  2.  The  central  apparatus  is  de- 
veloped in  response  to  and  especially  adapted  to  receive  the  specific  stimulus. 
The  interpretations  that  are  made  in  consciousness  in  response  to  an  inflow 
of  nerve  impulses  from  the  sense  organ  are  more  or  less  constant.  When 
the  exceptional  stimulus  is  applied  to  the  special  end  organ  it  results  in  the 
usual  change  in  the  sense  organ,  the  resulting  nerve  impulses  reach  the  usual 
central  area,  and  there  is  no  physiological  basis  for  other  than  the  usual  sen- 
sations which  development  and  experience  have  associated  with  activity  in 
the  parts  affected. 

Transmission  of  Nerve  Impulses  through  the  Neurone. — The  the- 
ory has  been  advanced  that  in  the  nerve  cell  the  primary  function  of  some 
processes  is  to  carry  nerve  impulses  toward  the  cell  body,  and  of  other  proc- 
esses to  carry  nerve  impulses  away  from  the  cell  body.  At  the  present  time 
this  view  is  advocated  by  perhaps  the  ablest  living  anatomists  and  neurolo- 
gists. The  dendrites  conduct  toward  the  cell  body,  and  the  axones  away  from 
it.  That  is,  the  former  are  cellulipetal,  the  latter  cellulifugal. 

Impressions  made  upon  the  terminations  or  upon  the  trunk  of  an  afferent 
nerve  may  cause,  a,  pain  or  some  other  kind  of  general  sensation;  fc,  special 
sensation;  c,  reflex  action  of  some  kind;  or  d,  inhibition  or  restraint  of  action. 
Similarly  impressions  made  upon  an  efferent  nerve  may  cause,  a,  contraction 
of  muscle  (motor  nerve);  b,  it  may  influence  secretion  (secretory  nerve); 
c,  it  may  influence  nutrition  (trophic  nerve) ;  or  d,  it  may  inhibit,  augment, 
or  stop  any  other  efferent  action. 

By  artificial  stimulation  nerve  impulses  can  be  made  to  pass  in  both  di- 
rections in  all  classes  of  nerve  processes.  That  is  to  say,  if  a  motor  axone  is 
artificially  stimulated  in  the  middle  of  its  course  it  will  not  only  convey  a  nerve 
impulse  to  its  distribution,  but  also  a  nerve  impulse  will  pass  back  over  the  fiber 
to  the  cell  body  and  out  over  the  dendrites.  Normally,  in  the  complex  of  the 
body,  it  is  probable  that  such  a  neurone  will  be  stimulated  only  at  its  points 
of  contact  with  other  neurones  chiefly  through  its  dendrites,  and  especially 
by  means  of  the  sensory  cells.  The  dendrites  therefore  will  receive  the  nerve 
stimulus,  carry  it  through  the  cell  body  to  the  axone  and  its  distribution. 
In  such  cells  there  is  isolated,  or  uninterrupted,  conduction  throughout  the 
extent  of  the  neurone.  The  nerve  impulse  is  able  to  pass  from  a  given 
neurone  to  adjacent  ones  only  at  the  termination  of  the  axone  or  its  branches, 

37 


578  THE    NERVOUS    SYSTEM 

and  such  terminations  may  be  considered  as  special  organs  for  the  trans- 
ference of  the  nerve  impulses.  This  activity  involves  isolated  conduction  in 
nerve  fibers  bound  in  a  common  nerve  trunk.  It  has  been  supposed  that  the 
myelin  sheath  of  a  medullated  nerve  acts  as  an  insulator  of  the  axis-cylinder, 
but  this  can  be  only  relatively  true,  for  the  reason  that  non-medullated  nerves 
do  not  possess  the  myelin  sheath.  In  non-medullated  nerves  we  must  sup- 
pose that  the  primitive  sheath  is  sufficient  to  give  insulated  conduction,  or 
that  it  is  an  inherent  property  of  the  axis-cylinder  itself  to  carry  the  nerve 
impulse  without  transmission  to  adjacent  fibers. 

We  have  already,  page  500,  discussed  the  rate  of  transmission  of  the  nerve 
impulse  in  motor  nerves  which  was  given  as  from  27  to  30  meters  per  second. 
In  sensory  nerves  the  rate  is  said  to  be  somewhat  higher;  in  human  nerve 
from  30  to  42  meters  per  second. 

Physiological  Types  of  Neurones. — Many  classifications  could  be 
made  of  nerve  cells,  based  on  the  differences  in  their  functional  relations 
but  attention  will  be  called  to  only  one.  Neurones  may  be  classified  as 
afferent  or  sensory,  efferent  or  motor,  and  connecting  or  transmitting. 

Under  afferent  neurones  are  classed  all  those  neurones  which  transmit 
the  effects  of  external  stimuli  received  through  the  sense  organs,  both  general 
and  special  sense  organs.  These  neurones  carry  nerve  impulses  toward  the 
central  nervous  system,  which  may  ultimately  produce  those  changes  in  the 
cerebral  cortex  which  are  associated  with  states  of  consciousness. 

Under  efferent  neurones  are  included  all  those  which  transmit  nerve  im- 
pulses from  any  part  of  the  central  nervous  system  to  the  muscles,  that  is, 
motor  nerves;  or  transmit  nerve  impulses  to  the  glands,  secretory  nerves;  or 
that  transmit  nerve  impulse,  which  inhibit  peripheral  action,  inhibitory 
nerves. 

Under  central  or  transmitting  neurones  are  included  those  units  which 
act  as  connecting  links  within  the  central  organ,  especially  within  coordi- 
nate parts  of  the  central  nervous  system,  between  the  afferent  and  efferent 
neurones. 

Nerve  Centers. — Whenever  a  number  of  neurones  are  gathered  in 
one  group  to  accomplish  some  specific  function  it  is  called  a  nerve  center. 
The  term  usually  applies  to  the  aggregation  of  cell  bodies  and  their  dendritic 
processes  in  contradistinction  to  nerve  trunks.  There  are  aggregations 
of  nerve  cells  into  different  specific  groups,  to  which  we  cannot  in  every 
case  ascribe  a  specific  function.  These  groups  are  not  called  nerve  centers, 
but  are  described  by  the  general  anatomical  term,  ganglia.  Such  ganglia 
are  represented  in  the  sympathetic  chain,  the  spinal-root  ganglia,  the  ganglia 
of  certain  cranial  nerves,  etc.  The  nerve  centers  are  found  in  the  spinal  cord, 
the  medulla,  and  the  higher  cranial  groups.  The  medulla  is  particularly 
rich  in  nerve  centers.  The  cerebro-spinal  axis  is  in  fact  an  aggregation  of 
nerve  centers  of  varying  complexity  in  different  parts. 


NERVE    CENTERS  579 

It  is  by  means  of  the  nerve  centers  that  the  activities  of  the  differentiated 
parts  of  the  human  body  are  brought  into  intimate  correlation.  The  nerve 
centers  exercise  their  influence  through  the  power  of  inhibiting  or  decreasing 
activity;  or,  on  the  other  hand,  of  augmenting  or  increasing  the  activity  in  the 


FIG.  359.— View  of  the  Cerebro-spinal  Axis  of  the  Nervous  System.  The  right  half  of 
the  cranium  and  trunk  of  the  body  has  been  removed  by  a  vertical  section;  the  mem- 
branes of  the  brain  and  spinal  cord  have  also  been  removed,  and  the  roots  and  first  part  of 
the  fifth  and  ninth  cranial,  and  of  all  spinal  nerves  of  the  right  side,  have  been  dissected 
out  and  laid  separately  on  the  wall  of  the  skull  and  on  the  several  vertebrae  opposite  to  the 
place  of  their  natural  exit  from  the  cranio-spinal  cavity.  (After  Bourgery.) 


580  THE    NERVOUS    SYSTEM 

peripheral  tissues  or  in  other  parts  of  the  nervous  system.  For  example,  the 
vagus  center  regulates  the  activity  of  the  heart  muscle  by  its  power  to  decrease 
or  inhibit  cardiac  contractions.  This  center,  we  have  already  found,  is  in 
constant  tonic  activity;  that  is  to  say,  in  constant  regulative  control  of  the 
heart.  The  cardiac  augmentory  center,  on  the  other  hand,  produces  just  the 
opposite  effect,  increasing  the  activity  of  the  cardiac  muscle.  What  is  true 
for  the  heart  is  likewise  true  in  general  for  other  tissues  of  the  body.  The 
numerous  nerve  centers  in  the  central  nervous  system  are  brought  into  cor- 
relation through  an  exceedingly  complex  system  of  communicating  fibers. 
The  cerebro-spinal  axis  may  in  fact  be  regarded  as  a  segmental  chain  of  nerve 
centers,  the  complexity  increasing  from  the  cord  toward  the  brain,  and  the 
coordinating  control  culminating  in  the  cerebral  cortex. 

THE  STRUCTURE  AND  FUNCTION  OF  THE 
SPINAL  CORD. 

STRUCTURE. 

The  spinal  cord  is  a  cylindrical  column  of  nerve-substance  connected 
above  with  the  brain  through  the  medium  of  the  bulb,  and  terminating  below 
in  a  slender  filament  of  nerve  substance,  the  filum  terminale,  which  lies  in  the 
midst  of  the  roots  of  the  many  nerves  forming  the  cauda  equina. 

General  Features. — The  cord  is  composed  of  nerve  fibers  and  nerve 
cells.  The  former  are  situated  externally  and  constitute  the  chief  portion 
of  the  cord,  while  the  latter  occupy  its  central  or  axial  portion  and  are  so 
disposed  that  on  the  surface  of  a  transverse  section  of  the  cord  two  somewhat 
crescentic  grayish  masses  connected  by  a  narrower  portion  or  isthmus  ap- 
pear, figure  358.  Passing  through  the  center  of  the  cord  in  a  longitudinal 
direction  is  a  minute  canal,  the  central  canal,  which  is  continued  through 
the  whole  length  of  the  cord,  opening  above  into  the  space  at  the  back  of 
the  medulla  oblongata  and  pons  Varolii  called  the  fourth  ventricle.  The 
canal  is  lined  by  a  layer  of  columnar  ciliated  epithelium. 

The  spinal  cord  consists  of  exactly  symmetrical  halves,  separated  an- 
teriorly and  posteriorly  by  vertical  fissures  (the  posterior  fissure  being  deeper 
but  less  wide  and  distinct  than  the  anterior),  and  united  in  the  middle  by 
nervous  matter  which  forms  the  commissures.  The  central  part,  which 
contains  the  central  canal,  is  known  as  the  gray  commissure,  and  is  bounded 
by  the  anterior  white  commissure  in  front.  Each  half  of  the  spinal  cord  is 
marked  on  the  sides  (obscurely  at  the  lower  part,  but  distinctly  above)  by 
two  longitudinal  furrows,  which  divide  it  into  three  portions,  funiculi,  or  tracts 
— an  anterior,  lateral,  and  posterior.  From  the  groove  between  the  anterior 
and  lateral  funiculi  spring  the  anterior  roots  of  the  spinal  nerves;  and  just 


GENERAL   FEATURES  50 1 

in  front  of  the  groove  between  the  lateral  and  posterior  funiculi  arise  the 
posterior  roots  of  the  same;  a  pair  of  roots  on  each  side  corresponding  to 
each  segment  of  the  cord. 

The  nerve  tracts  of  the  cord  are  made  up  of  medullated  nerve  fibers  of 
different  sizes,  arranged  longitudinally,  and  of  a  supporting  material  of 
ordinary  fibrous  connective  tissue  and  neuroglia,  figure  105. 


IS    15 


16         17 


13- 


f 
I 

if 


FIG.  360. — Horizontal  Section  of  the  Cord  and  its  Envelopes,  at  the  Middle  of  a 
Vertebral  Body  (Schematic),  i,  Spinal  cord  with  2,  its  anterior  median  fissure;  3,  its 
posterior  median  fissure;  4,  anterior  roots;  5,  posterior  roots;  6,  pia  mater  (in  red);  7, 
ligamentum  dentatum;  8,  connecting  fibers  passing  from  the  pia  to  dura  mater;  9,  visceral 
layer,  and  9',  parietal  layer  of  the  arachnoid  (in  blue);  10,  subarachnoid  space;  n,  arach- 
noid cavity;  12,  dura  mater  (in  yellow);  13,  periosteum;  13',  external  periosteum;  14, 
cellular  tissue  situated  between  the  dura  mater  and  the  wall  of  the  vertebral  canal;  15, 
common  posterior  vertebral  ligament;  16,  intraspinal  veins;  17,  vertebra  in  section. 
(Testut.) 


The  general  rule  respecting  the  size  of  different  segments  of  the  cord 
appears  to  be  that  each  is  in  direct  proportion  in  this  respect  to  the  size  and 
number  of  nerve  roots  given  off  from  it,  and  has  but  little  relation  to  the  size 
or  number  of  those  given  off  below  it.  Thus  the  cord  is  very  large  in  the 
middle  and  lower  part  of  its  cervical  portion,  whence  arise  the  large  nerve 
roots  for  the  formation  of  the  brachial  plexuses  and  the  nerve  supply  of  the 
upper  extremities;  and  again  enlarges  at  the  lowest  part  of  its  dorsal  portion 
and  the  upper  part  of  its  lumbar,  at  the  origins  of  the  large  nerves  which, 


582 


THE    NERVOUS    SYSTEM 


after  forming  the  lumbar  and  sacral  plexuses,  are  distributed  to  the  lower 
extremities.  The  chief  cause  of  the  greater  size  at  these  parts  of  the  spinal 
cord  is  increase  in  the  quantity  of  the  gray  matter;  for  there  seems  reason 
to  believe  that  the  white  part  of  the  cord  becomes  gradually  and  progressively 
larger  from  below  upward,  doubtless  from  the  addition  of  a  certain  number 
of  ascending  fibers  from  each  pair  of  nerves. 

From  careful  estimates  of  the  number  of  nerve  fibers  in  a  transverse  sec- 
tion of  the  cord  toward  its  upper  end,  and  the  number  entering  or  issuing 
from  it  by  the  anterior  and  posterior  roots  of  each  pair  of  nerves,  it  has  been 
shown  that  in  the  human  spinal  cord  not  more  than  half  of  the  total  number 


FIG.  361. — From  the  Lower  Lumbar  Cord  of  Man,  after  a  Preparation  by  Klonne  and 
Muller  stained  by  Weigert  and  Pal's  method.  A  portion  of  the  gray  substance  of  the 
anterior  column  with  the  adjoining  portions  of  the  lateral  funiculus  is  represented,  showing 
anterior  column  cells  and  the  fine  medullated  fibers  which  enter  the  gray  substance  from  the 
lateral  funiculi  and  surround  the  nerve  cells,  which  here  are  provided  with  fine  pigmented 
granules.  High  power.  (Kolliker.) 

of  nerve  fibers  of  all  the  spinal  nerves  are  contained  in  a  transverse  section 
near  its  upper  end.  It  is  obvious,  therefore,  that  at  least  half  of  the  nerve 
fibers  entering  it  must  terminate  somewhere  within  the  cord  itself. 

The  Arrangement  of  Nerve  Cells  in  the  Spinal  Cord. — The  gray  mat- 
ter of  the  spinal  cord  consists  of  numerous  groups  of  nerve  cells  and  of  a 
close  mesh  work  of  nerve  fibers,  most  of  which  are  very  fine  and  delicate. 


ARRANGEMENT    OF   NERVE    CELLS    IN    SPINAL   CORD  583 

Medullated  fibers  mingled  with  the  small  gray  fibers  about  the  borders  of  the 
gray  substance.  Mingled  with  it  and  supporting  it  is  the  meshwork  of  the 
neuroglia. 

The  multipolar  cells  of  the  cord  are  either  scattered  singly  or  arranged  in 
groups  or  columns  in  bilateral  symmetry.  The  following  are  to  be  distin- 
guished, certain  of  the  groups  being  more  or  less  marked  in  all  of  the  regions 
of  the  cord,  viz.,  a,  those  in  the  anterior  columns;  b,  those  in  the  lateral 
columns;  c,  those  in  the  posterior  columns;  and  d,  intrinsic  cells  distributed 
throughout  the  gray  matter. 


FIG.  362. — Section  of  Spinal  Cord,  One  Half  of  Which  (Left)  Shows  the  Tracts  of  the 
White  Matter,  and  the  Other  Half  (Right)  Shows  the  Position  of  the  Nerve  Cells  in  the 
Gray  Matter.  7,  10,  9,  and  3  are  tracts  of  descending  degeneration;  i,  2,  4,  6,  and  8,  of 
ascending  degeneration.  Semidiagrammatic.  See  the  text  for  a  description  of  the  groups 
of  nerve  cells  shown  on  the  right.  (After  Sherrington.) 

The  cells  in  the  anterior  columns  are  large  and  branching,  and  each  gives 
rise  to  an  axis-cylinder  process  which  passes  out  in  the  anterior  nerve  root. 
These  cells  are  everywhere  conspicuous,  but  are  particularly  numerous 
in  the  cervical  and  lumbar  enlargements.  In  these  districts  they  may  be 
divided  into  several  groups;  i.  A  group  of  large  cells  close  to  the  tip  of  the 
inner  part  of  the  anterior  column,  i,  figure  362.  This  group  is  called  the 
antero-mesial  group  of  motor  cells.  It  forms  r,  column  the  full  length  of  the 
cord  and  is  supposed  to  furnish  motor  innervation  to  the  long  muscles  of  the 
trunk.  2.  The  antero-lateral  group  of  cells,  2a,  forms  a  column  which  has 
its  best  development  in  the  cervical  and  lumbo-sacral  enlargements.  It 
probably  furnishes  motor  fibers  to  the  muscles  of  the  limbs  including  those 
of  the  shoulders  and  hips.  3.  There  are  groups,  2b,  2C,  the  intermedio  and 
lateral  groups,  which  form  slender  columns  in  the  cord,  through  the  entire 


584  THE    NERVOUS    SYSTEM 

dorsal  and  the  first  two  lumbar  segments.  The  cells  are  small  and  closely 
aggregated  and  characteristic  in  appearance.  These  nuclei  reappear  in  the 
upper  cervical  segments  and  in  the  sacral  segments  that  give  origin  to  the 
nervi  erigens.  They  are  supposed  to  be  the  motor  nuclei  for  the  vaso-motor, 
the  pilo-motor  nerves,  and  for  the  nerves  of  the  sweat  glands,  i.e.,  the  auto- 
nomic  system.  4.  The  cells  of  the  posterior  vesicular  column,  or  Clark's 
column,  are  in  the  posterior  portion  of  the  gray  matter  of  the  cord.  They 
form  a  conspicuous  column  of  cells,  3,  5,  and  6  of  figure  362,  extending  from 
the  last  cervical  to  the  second  lumbar  segments.  These  large  cells  contribute 
their  axones  to  the  ascending  cerebello-spinal  (direct  cerebellar)  fasciculus 
which  ascends  to  the  cerebellum  and  is  believed  to  carry  sensory  impulses  of 
prime  importance  in  the  coordinations  of  muscular  movements.  The  nerve 
relations  and  the  restricted  location  of  the  posterior  vesicular  column  has  also 
led  to  the  suggestion  that  its  function  has  to  do  with  the  visceral  afferent  or 
sensory  nerve  impulses.  Sensory  fibers  from  the  posterior  roots  arborize 
around  the  cells  of  the  posterior  vesicular  column,  i.e.,  the  dorsal  nucleus,  see 
figure  363.  The  dorsal  nucleus  is  considerably  broken  up  by  the  passage  of 
bundles  of  fibers  through  it,  called  the  lateral  reticular  formation. 

Besides  these  groups,  which  have  their  names  largely  on  account  of  their 
location,  there  are  distributed  throughout  the  gray  matter  a  very  large  num- 
ber of  other  cells,  which  are  known  as  intrinsic  cells.  These  are  of  two  types, 
those  restricted  to  connections  wholly  within  the  gray  matter  Golgi  cells  of 
the  second  type  and  those  that  send  out  axones  which  pass  into  the  adjacent 
ground  bundles  of  the  same  or  of  the  opposite  side,  and  pass  up  and  down  the 
cord,  to  enter  the  gray  matter  again.  They  connect  by  their  end-brushes  with 
cells  at  different  levels  of  the  cord. 

The  functions  of  these  connecting  or  intrinsic  cells  is  to  unite  the  pos- 
terior and  anterior  regions  of  the  cord,  to  serve  as  conductors  between  the 
lateral  halves,  or  to  connect  segments  at  different  levels.  They  are  also  dis- 
tributing fibers  in  that  they  bring  a  single  or  at  least  a  small  number  of  pos- 
terior sensory  neurones  into  connection  with  a  relatively  large  number  of 
anterior  or  motor  neurones. 

The  Fasciculi  or  Tracts  in  the  White  Matter  of  the  Spinal  Cord.— 
We  have  already  shown  that  the  white  matter  of  the  cord  is  divided  on  each 
side  into  the  anterior,  lateral,  and  posterior  funiculi.  But  evidence  of  various 
kinds  discussed  below  has  shown  the  following  main  conducting  channels  or 
tracts  through  the  cord,  see  figures  363  and  364  for  illustration. 

Tracts  of  the  Cord. 

Funiculus  posterior. 

Fasciculus  gracilis,  tract  of  Goll  (ascending). 
Fasciculus  cuneatus,  tract  of  Burdach  (ascending) 
Comma,  tract  of  Schultz  (descending). 


FASCICULI  OR  TRACTS  IN  WHITE  MATTER  OF  SPINAL  CORD       585 

Funiculus  lateralis. 

Fasciculus  cerebello-spinalis,  direct  cerebellar  (ascending). 
Fasciculus  antero-lateralis  super  ficialis,  tract  of  Gowers  (ascending). 
Fasciculus    cerebro-spinalis    lateralis,    crossed    pyramidal    tract 
(descending). 


Radix  anterior 
Fasciculus  cerebrospinalis  lateralis  fpyramidalis  lateralisj    4 

Anterior  root  fibre 

Bundle  to  anterior  funtculus  from  the  formatio  reticularis     _ 

Fasciculus  cerebrospinalis  anterior 

[pyramidalis  anterior*] 


\ 


Bundle  to  anterior  funiculus    , 
from  the  fprmatio  reticularis  \ 


Substantia  grisea.  — 


Fasciculus 

anterolateral.is  supcr- 

ftcialis  [Gowersi]  and 

(ascending)  bundle  from 

anterior  funiculus  to  the 

formatio  reticularis 

Substantia  alba 

Bundte  to  lateral 
funiculus  from  Betters' 
nucleus  and  from  "tho 
red  nucleus . 


Fasciculus  cferebro 
splnalis  lateralis 
lateral! 


Nervus  spinalis 

y  Ganglion  spinal e 

Cells  of  tho  spinal  ganglion 
Fasciculus  cerebellospinali* 
^  Secondary  reflex  path 

f  Radix  posterior 

Collateral 

to  the  posterior  horn 

Secondary  path  of 

posterior  funiculus 

.  fijescending  posterior 

root  fibre 
•  Primary  reflex  -path 

Ascending  posterior 
root  fibre 


''.->  Secondary  reflex  patU 


.Descending  posterior 
root  fibre 


—  Posterior  foot  fibre 


Sulcus  medianus 
posterior 


—  Posterior  root  fibre 


"FiG.  363. — Reconstruction  of  a  Segment  of  the  Spinal  Cord  Representing  Both  a  Trans- 
verse and  Longitudinal  Section.     (Held,  from  Spalteholz's  Anatomy.) 


Fasciculus  lateralis  proprius,  lateral  ground  bundles  (ascending 
and  descending). 

Funiculus  anterior. 

Fasciculus  cerebro-spinalis  anterior,  direct  pyramidal  tract   (de- 
scending). 

Fasciculus  anterior  proprius,  anterior  ground  bundles  (ascending 
and  descending). 


586  THE    NERVOUS    SYSTEM 

The  methods  for  determining  the  tracts  in  the  cord  and  the  evidence  on 
basis  of  which  function  is  ascribed  are  briefly  the  following: 

The  Embryological  Method. — It  has  been  found  that,  if  the  development 
of  the  spinal  cord  be  carefully  observed  at  different  stages,  certain  groups  of 
nerve  fibers  acquire  their  myelin  sheath  at  earlier  periods  than  others,  and 
that  the  different  groups  of  fibers  can  therefore  be  traced  in  various  directions. 
This  is  known  as  the  method  of  Flechsig. 

Wallerian  or  Degeneration  Method. — This  method  depends  upon  the  fact 
already  presented  that  if  a  nerve  fiber  is  separated  from  its  nerve  cell  it 
degenerates.  It  consists  in  tracing  the  course  of  tracts  of  degenerated  fibers 
which  result  from  an  injury,  or  lesions  in  any  part  of  the  central  nervous 
system.  When  fibers  degenerate  below  a  lesion  the  tract  is  said  to  be  of 
descending  degeneration,  and  when  the  fibers  degenerate  in  the  opposite 
direction  the  tract  is  one  of  ascending  degeneration.  By  modern  methods  of 
staining  of  the  nervous  tissue,  it  has  proved  comparatively  easy  to  distinguish 
degenerated  parts  in  sections  of  the  cord  or  of  other  portions  of  the  central 
nervous  system.  Degenerated  fibers  have  a  differential  staining  reaction 
when  the  sections  are  treated  by  Marchi's  method.  Accidents  resulting  in 
loss  of  function  and  in  degeneration  in  the  central  nervous  system  in  man 
have  given  us  much  information  as  to  its  organization,  but  this  has  of  late 
years  been  supplemented  and  largely  extended  by  the  experiments  on  animals, 
particularly  upon  monkeys.  Considerable  light  has,  by  the  method  of 
section  and  degeneration,  been  shed  upon  the  path  of  conduction  of  impulses 
to  and  from  the  various  parts  of  the  nervous  system.  Thus  we  not  only 
have  embryological  evidence  mapping  out  different  tracts,  but  also  con- 
firmatory pathological  and  experimental  observations. 

T'he  tracts  which  have  been  made  out  are  the  following: 

Tracts  of  Descending  Degeneration. — The  Lateral  Pyramidal  Tract.— 
This  tract  is  situated  to  the  outer  side  of  the  posterior  cornu  of  gray  matter, 
figure  360,  7.  It  originates  in  the  cerebral  cortex  and  extends  throughout 
the  whole  length  of  the  spinal  cord;  at  the  lower  part  it  extends  to  the  margin 
of  the  cord,  but  higher  up  it  becomes  displaced  inward  from  this  position  by 
the  interpolation  of  another  tract  of  fibers,  the  direct  cerebellar  tract.  The 
lateral  pyramidal  tract  is  large,  and  may  touch  the  tip  of  the  gray  matter  of 
the  posterior  gray  column,  but  it  is  separated  from  it  elsewhere.  It  is  oval 
in  shape  on  cross- section,  and  diminishes  in  size  from  the  cervical  region 
downward.  The  tract  is  particularly  well  marked  out,  both  by  the  degener- 
ation and  by  embryological  methods.  The  fibers  are  supposed  to  pass  off 
as  they  descend,  and  to  join  the  various  local  nervous  mechanisms  of  nerve 
cells  and  their  branchings  which  are  represented  in  the  anterior  columns  of 
the  cord.  The  fibers  of  which  this  tract  is  composed  are  moderately  large, 
but  are  mixed  with  some  that  are  smaller. 

The  Anterior  Pyramidal  Tract. — This  tract  is  situated  in  the  anterior 


THE    SPINAL   NERVES  587 

funiculus  by  the  sides  of  the  anterior  fissure,  figure  360,  10.  It  is  smaller 
than  the  lateral  tract  and  is  not  present  in  all  animals,  though  conspicuous 
in  the  human  cord  and  in  that  of  the  monkey.  It  can  be  traced  upward  to 
the  cerebral  cortex,  and  downward  as  far  as  the  mid  or  lower  thoracic  region, 
where  it  ends. 

Antero-lateral  Descending  Tract. — This  is  an  extensive  tract,  elon- 
gated but  narrow,  and  reaching  from  the  lateral  pyramidal  to  the  anterior 
pyramidal  tract.  It  is  a  mixed  tract,  since  not  all  of  its  fibers  degenerate 
below  the  lesions. 

Comma  Tract. — This  is  a  small  tract  of  fibers  in  the  posterior  funiculus 
which  degenerates  below  the  point  of  section  of  the  cord.  It  consists  of  the 
descending  collaterals  of  the  posterior  nerve  roots  as  they  pass  into  the 
cord. 

Tracts  of  Ascending  Degeneration. — The  fasciculus  gracilis,  tract  of 
Goll,  and  the  fasciculus  cuneatus,  tract  of  Burdach.  These  tracts  degenerate 
upward  on  section  of  the  cord,  also  on  section  of  the  posterior  nerve  roots, 
figure  362,  i.  They  exist  throughout  the  whole  of  the  cord  and  can  be  traced 
into  the  bulb.  They  are  sensory  tracts,  see  page  586. 

The  Fasciculus  Cerebello-spinalis,  or  Direct  Cerebellar  Tract. — This 
tract  is  situated  on  the  outer  part  of  the  cord  between  the  lateral  pyramidal 
tract  and  the  margin.  It  is  found  in  the  cervical,  thoracic,  and  uppjf 
lumbar  regions  of  the  cord,  and  increases  in  size  from  below  upward.  It 
degenerates  on  injury  or  section  of  the  cord  itself,  but  not  on  section  of  the 
posterior  nerve  roots,  since  its  fibers  arise  from  the  cells  of  the  dorsal  nucleus. 
As  the  name  implies,  it  is  believed  to  pass  up  into  the  cerebellum,  see 
Page  597. 

The  Fasciculus  Antero-lateralis,  Tract  of  Gowers. — This  tract  lies 
at  the  margin  of  the  lateral  funiculus  of  the  cord  and  extends  the  full  length, 
see  figure  364. 

It  will  thus  be  seen  that  the  three  general  divisions  of  the  white  matter 
of  the  spinal  cord — the  anterior,  the  lateral,  and  posterior  funiculi — are 
subdivided  into  tracts  in  which  the  fibers  degenerate  upward,  those  in  which 
the  fibers  degenerate  downward,  and  still  others  in  which  the  fibers  degener- 
ate either  way  for  only  short  distances  when  the  cord  is  cut  across.  These 
latter  parts  of  the  cord  are  composed  of  association  fibers  which  connect 
different  levels  of  the  cord.  The  association  tracts  form  the  antero-lateral 
columns  and  the  lateral  limiting  layer.  The  arrangement  of  these  tracts  is 
shown  well  in  figures  362  and  364. 

The  Spinal  Nerves. — The  spinal  nerves  consist  of  thirty-one  pairs, 
from  the  sides  of  the  whole  length  of  the  cord,  their  number  corresponding 
with  the  intervertebral  foramina  through  which  they  pass.  Each  nerve 
arises  by  two  roots,  an  anterior  and  a  posterior,  the  latter  being  the  larger. 
The  roots  emerge  through  separate  apertures  of  the  sheath  of  dura  mater 


588 


THE    NERVOUS    SYSTEM 


surrounding  the  cord;  and  directly  after  their  emergence,  where  the  roots  lie 
in  the  intervertebral  foramen,  a  ganglion  is  found  on  the  posterior  root.  The 
anterior  root  lies  in  contact  with  the  anterior  surface  of  the  ganglion,  but  none 
of  its  fibers  intermingle  with  those  in  the  ganglion,  figure  361.  But  imme- 
diately beyond  the  ganglion  the  two  roots  coalesce,  and  by  the  mingling  of 
their  fibers  form  a  mixed  spinal  nerve;  the  spinal  nerve,  after  issuing  from 
the  intervertebral  canal,  gives  off  anterior  and  posterior  (or  ventral  and  dorsal) 
branches,  each  containing  fibers  from  both  the  roots.  A  third  or  visceral 
branch  of  the  spinal  nerve,  ramus  communicans,  joins  the  sympathetic 
chain. 

The  anterior  root  of  each  spinal  nerve  arises  by  numerous  separate  and 


Entering  posterior 
root 


Lissauer's  tract 


anterior  root 


FIG.  364. — Diagrammatic  Transverse  Section  of  the  Spinal  Cord,  Showing  the  Conduction 
Paths  and  Groups  of  Cells.     (Cunningham.) 


converging  bundles  from  the  anterior  columns  of  the  cord;  the  posterior  root 
by  more  numerous  parallel  bundles,  from  the  posterior  column.  The 
anterior  roots  of  each  spinal  nerve  consist  chiefly  of  efferent  fibers;  the  pos- 
terior exclusively  of  afferent  fibers. 

Course  of  the  Fibers  of  the  Spinal  Nerve  Roots.— The  Anterior 
Roots. — The  anterior  roots  leave  the  cord  in  several  bundles,  which  may  be 
called:  i,  Internal;  2,  Middle;  3,  External.  All  have  their  origin  from  the 
groups  of  multipolar  cells  in  the  anterior  columns.  The  internal  fibers  are 
originated  partly  in  the  internal  group  of  nerve  cells  of  the  anterior  columns 


COURSE    OF    THE    FIBERS    OF    THE    SPINAL   NERVE    ROOTS         589 

of  the  same  side;  but  a  few  fibers  can  be  traced  through  the  anterior  com- 
missure to  cells  of  the  anterior  column  of  the  opposite  side. 

The  Posterior  Roots. — The  fibers  of  the  posterior  roots  enter  the  spinal 
cord  to  the  inner  or  median  side  of  the  posterior  column.  The  fibers,  as 
soon  as  they  reach  the  cord,  divide  in  a  fork-like  fashion,  one  branch  pass- 
ing down  a  short  distance,  about  three  centimeters,  the  other  branch  passing 
up  for  a  shorter  or  longer  distance.  This  upper  branch  sometimes  reaches 
the  whole  extent  of  the  cord,  but  generally  it  extends  over  only  one  or  two 
segments  of  the  cord.  The  divisions  of  the  posterior  root  fibers  give  off 


FIG.  365. — Section  of  the  Spinal  Cord,  Showing  the  Grouping  of  Nerve  Cells  and  the 
Course  of  Nerve  Fibers  Entering  in  Posterior  and  Anterior  Roots.  Numerals  1-6  are 
different  types  of  the  sensory  fibers.  Their  collaterals,  7-1 2,  connect  with  different  regions 
of  the  gray  substance;  13  and  14,  collaterals  from  descending  tracts  of  the  cord;  r.a., 
anterior  root;  a,  anterior  motor  cells;  6,  cells  contributing  fibers  to  the  various  tracts; 
c,  commissural  cells;  d,  Golgi  cells.  (After  Lenhossek.) 


in  their  course  numerous  collaterals,  figure  368.  The  fibers  of  the  posterior 
roots  are  divided  into  two  sets,  an  internal  or  median,  an  external  or  lateral 
group.  The  lateral  set  consists  mostly  of  small  fibers  which  enter  the 
cord  opposite  the  tip  of  the  posterior  horn.  The  fibers  pass  in  part  to  the 
marginal  column  of  Lissauer,  where  they  ascend  and  descend;  in  part  they 
penetrate  the  posterior  column,  and  come  in  relation  with  its  cells.  From 
the  median  set  some  fibers  pass  to  Clarke's  column,  others  pass  by  way 
of  the  posterior  commissure  to  the  median  cells  of  the  other  side.  Others 
pass  through  the  gray  matter  to  the  anterior  column  cells  of  the  same  side, 


5 QO  THE    NERVOUS    SYSTEM 

figure  365.  Besides  this,  they  are  connected  through  collaterals  with  the 
intrinsic  cells  of  the  gray  matter  at  different  levels  of  the  cord.  One  can 
realize  that  each  nerve  root  has,  in  this  way,  an  effective  grip  upon  a  large 
extent  of  the  cord.  This  is  seen  well  by  studying  figures  363  and  365. 

The  Peculiarities  of  Different  Regions  of  the  Spinal  Cord.— The 
outline  of  the  gray  matter  and  the  relative  proportion  of  the  white  matter 
vary  in  different  regions  of  the  spinal  cord,  and  it  is,  therefore,  possible  to 
tell  approximately  from  what  region  any  given  transverse  section  of  the 
spinal  cord  has  been  taken.  The  white  matter  increases  in  amount  from 
below  upward.  The  amount  of  gray  matter  varies;  it  is  greatest  in  the 
cervical  and  lumbar  enlargements,  viz.,  at  and  about  the  5th  lumbar  and  the 
6th  cervical  nerves,  and  least  in  the  thoracic  region.  The  greatest  develop- 
ment of  gray  matter  corresponds  with  greatest  number  of  nerve  fibers  pass- 
ing from  the  cord. 

In  the  cervical  enlargement  the  gray  matter  occupies  a  large  proportion  of 
the  section,  the  gray  commissure  is  short  and  thick,  the  anterior  column  is 
blunt,  while  the  posterior  is  somewhat  tapering.  The  anterior  and  posterior 
roots  run  some  distance  through  the  white  matter  before  they  reach  the 
periphery. 

In  the  dorsal  region  the  gray  matter  bears  only  a  small  proportion  to  the 
white,  and  the  posterior  roots  in  particular  run  a  long  course  through  the 
white  matter  before  they  leave  the  cord;  the  gray  commissure  is  thinner  and 
narrower  than  in  the  cervical  region. 

In  the  lumbar  enlargement  the  gray  matter  again  bears  a  very  large  propor- 
tion to  the  whole  size  of  the  transverse  section,  but  its  posterior  columns  are 
shorter  and  blunter  than  they  are  in  the  cervical  region.  The  gray  commis- 
sure is  short  and  extremely  narrow. 

At  the  upper  part  of  the  conus  medullaris,  which  is  the  portion  of  the  cord 
immediately  below  the  lumbar  enlargement,  the  gray  substance  occupies 
nearly  the  whole  of  the  transverse  section,  as  it  is  invested  only  by  a  thin  layer 
of  white  substance.  This  thin  layer  is  wanting  in  the  neighborhood  of  the 
posterior  nerve  roots.  The  gray  commissure  is  relatively  thick. 

At  the  level  of  the  -fifth  sacral  vertebra  the  gray  matter  is  again  in  excess,  and 
the  central  canal  is  enlarged,  appearing  T-shaped  in  section;  while  in  the 
upper  portion  of  the  filum  terminate  the  gray  is  uniform  in  shape  without  any 
central  canal. 

THE  FUNCTIONS  OF  THE  CORD. 

The  Reflex  Arc  and  Reflex  Action. — The  spinal  cord  is  morpho- 
logically a  segmental  or  metameric  structure.  This  is  shown  both  by  its 
development  and  by  its  comparative  anatomy.  The  pairs  of  nerves  are 
indicative  of  the  component  segments  of  the  cord.  The  tracts  of  the  cord  are 


THE  FUNCTIONS  OF  THE  CORD 


591 


in  a  sense  connectives  from  segment  to  segment,  connecting  the  cells  of 
both  adjacent  and  of  widely  separated  segments.     The  function  of  the  cord 


FIG.  366. — Schematic  Sketch  of  a  Reflex  Arc.  A ,  With  two  neurones,  an  afferent  and 
an  efferent;  B,  with  three  neurones,  an  afferent,  efferent,  and  a  connecting  or  intra- 
central  neurone. 


is  comprised  in  the  function  of  the  segments  and  in  the  function  of  the 
tracts. 

From  a  physiological  point  of  view,  it  may  almost  be  considered  as  an 
axiom  that  before  a  nerve  cell  can  send  out  a  nerve  impulse  it  must  first 
receive  a  stimulus  of  some  kind.  This  stimulus  usually  consists  of  an  afferent 
impulse  from  the  periphery.  Its  effect 
upon  the  receiving  cell  may  be  insufficient 
to  cause  any  response,  or  the  response  may 
be  delayed  for  a  long  period  and  may  in- 
volve many  complicated  nervous  activities 
and  even  psychological  processes.  Where 
the  peripheral  response  is  approximately 
immediate,  the  reaction  is  known  as  a 
reflex. 

A  reflex  arc,  reduced  to  its  simplest 
terms,  consists  of  the  following  anatomical 
elements:  a,  a  sensory  surface;  6,  an  affer- 
ent neurone;  c,  an  efferent  neurone;  J,  a 
muscle  or  gland.  The  simplest  form  of 
reflex  arc  is  schematically  shown  in  figures 
366  and  367. 

The  interlocking  of  fibers  between 
the  termination  of  the  afferent  neurone 
and  the  dendrite  of  the  efferent  neurone  FIG.  367.— Showing  the  Arrange- 

shown  in  figure   366  is  called  a  synapse.     ment  of  »  Simple  Reflex  Mechanism 

Composed  of  a  Motor  and  Sensor) 
1  he    reflex    arc   is   probably   seldom    as     Neurone,    sg,  Posterior  spinal  gan- 

simple    as    that    shown    in    figure     367,     Slion;  5  and  sth>  sensory  root;  m> 

motor-nerve  cell;   mw,   motor  root 
where   only    two   neurones   are  involved.     (Kolliker.) 


771 W, 


592 


THE    NERVOUS    SYSTEM 


More  often,  three  or  more  neurones  take  part,  as  shown  in  figures  366  B, 
and  368. 

The  neurone  connecting  the  afferent  neurone  with  the  efferent  neurone 
belongs  to  the  class  of  intracentral  or  connecting  neurones.  Since  all  parts 
of  the  cord,  in  fact  of  the  entire  cerebro-spinal  axis,  are  directly  or  indirectly 
associated  with  one  another  by  central  neurones,  figure  363,  the  possibility 
of  increasing  the  number  of  efferent  limbs  of  the  reflex  arc  can  be  readily 
understood. 

An  involuntary  physiological  reaction  in  a  tissue  produced  by  efferent 
nerve  impulses  which  have  been  discharged  from  a  nerve  center  under  the 

stimulus  of  a  sensory  or  afferent  nerve  im- 
pulse, is  called  a  reflex  act.  Where  the 
nervous  apparatus  involved  is  of  the  type 
represented  in  figures  366  and  367,  the 
activity  is  called  a  simple  reflex.  Most  re- 
flexes are  more  complex  in  character.  The 
afferent  nerve  impulse  passes  through  more 
than  one  simple  channel  in  the  cord,  so  that 
in  response  to  a  sensory  stimulus  a  series  of 
coordinated  acts  occurs  in  what  may  be 
called  a  complex  reflex. 

The  transmission  of  impulses  within  the 
cord  occurs  over  the  pathways  of  least  re- 
sistance. Increasing  a  number  of  synapses, 
or  the  number  of  neurone  links  in  the  chain 
of  conduction,  increases  the  resistance  so  that 

reflexes  will  occur  most  readily,  other  conditions  being  equal,  where  the  small- 
est number  of  neurones  is  involved,  i.e.,  in  the  same  segment  of  the  cord  in 
which  the  sensory  impulse  enters,  or  in  immediately  adjacent  segments.  In 
addition  to  the  number  of  synapses  in  the  reflex  arc,  certain  strictly  physio- 
logical factors  are  of  importance  in  determining  reflex  reaction;  e.g.,  the  in- 
tensity of  the  exciting  stimulus;  the  quality  of  the  stimulus;  the  rapidity  of 
the  recurrence  of  the  stimulus;  and  the  duration  of  its  application.  Thus, 
a  strong  stimulus  will  bring  about  a  reflex  reaction  sooner  than  a  weak  stimu- 
lus of  the  same  kind.  A  single  weak  stimulus  which  will  cause  no  reflex  may 
do  so  if  often  enough  and  rapidly  enough  repeated,  the  phenomenon  of 
summation  of  stimuli. 

A  reflex  act  once  started  may  result  in  efferent  impulses  which  continue 
for  some  time  after  the  exciting  cause  has  been  removed.  The  same  phe- 
nomenon is  observed  where  groups  of  nerve  cells  are  stimulated  directly.  It 
has  been  found,  by  observing  electrical  changes  in  nerve  fibers  by  means  of 
the  capillary  electrometer,  that  when  their  cells  of  origin  are  stimulated  they 
discharge  impulses  in  a  rhythmical  manner. 


FIG.  368. — Showing  the  Arrange- 
ment of  the  reflex  mechanism,  with 
a  neurone  intercalated  between  the 
sensory  and  motor  neurones. 


IRRADIATION    OF   IMPULSES    WITHIN    THE    CORD 


593 


Usually,  impulses  are  transmitted  to  a  nerve  cell  only  over  its  dendrite, 
but  it  must  be  also  assumed  that  such  a  conveyance  of  impulses  may  take 
place  over  the  collaterals  of  its  axone  near  the 
cell  body,  or  the  cell  body  may  be  stimulated 
directly  by  the  afferent  neurone.  The  per- 
ipheral fiber  of  the  spinal  ganglion  cell, 
although  it  has  the  structure  of  an  axone,  may 
be  looked  upon  physiologically  as  a  dendrite, 
since  homologues  in  lower  vertebrates  and 
in  man  himself  (nerve  cells  of  the  ganglion 
of  the  cochlea  of  the  auditory  nerve)  have 
this  structure,  the  nerve  cell  body  being 
situated  near  the  sensory  surface  from  which 
impressions  are  received. 

Irradiation  of  Impulses  within  the 
Cord. — Taking  as  an  example  a  frog  whose 
brain  has  been  destroyed,  a  simple  reflex  may 
be  demonstrated  by  irritating  the  skin  of  one 
foot  with  a  weak  stimulus.  In  response  to 
such  a  stimulus  the  foot  is  flexed  upon  the  leg, 
due  to  a  contraction  of  the  muscles  of  the 
reflex  arc  corresponding  to  the  sensory  surface 
irritated.  If  the  strength  or  duration  of  the 
stimulus  be  increased,  other  groups  of  muscles 
are  involved  in  the  following  order:  i.  Those 
of  the  leg  and  thigh  of  the  same  side;  2, 
homologous  muscles  of  the  opposite  side;  3, 
the  arms  of  the  same  side  and  of  the  opposite 
side. 

The  increasing  complexity  of  the  reflexes 
aroused  by  stimulation  of  one  and  the  same 
sensory  spot  is  not  easy  of  explanation.  We 
know  that  there  is  almost  an  infinite  number 
of  morphological  paths  in  the  cord,  yet  the 
responses  are  orderly  and  observe  a  certain 
sequence  in  their  increasing  complexity.  The 
reflexes  have  a  mechanical  definiteness  which, 
in  a  living  structure,  seems  almost  purposeful, 
yet  there  is  no  conscious  action  in  a  frog 
which  has  its  brain  destroyed. 

The  fact  is  that  in  the  development  of  the 
nervous  system  certain  physiological  paths  of  slight  resistance  have  been 

established  between  the  sensory  areas  and  the  muscles  which  move  the  parts 

38 


FIG.  369. — Scheme  of  Lower 
Motor  Neurone.  The  cell  body, 
protoplasmic  processes,  axone, 
collaterals,  and  terminal  arbor- 
izations in  muscle  are  all  seen  to 
be  parts  of  a  single  cell  and 
together  constitute  the  neurone. 
(Barker.)  c,  Cytoplasm  of  cell 
body  containing  chromophilic 
bodies,  neurofibrils,  and  peri- 
fibrillar  substance;  n,  nucleus;  n', 
nucleolus;  d,  dendrites;  ah,  axone 
hill  free  from  chromophilic  bodies; 
ax,  axone;  sf,  side  fibril  (col- 
lateral); m,  medullary  sheath;  nR, 
node  of  Ranvier  where  side 
branch  is  given  off;  si,  neuri- 
lemma  and  incisures  of  Schmidt; 
m,  striated  muscle  fiber;  tel, 
motor  end  plate. 


594  THE    NERVOUS    SYSTEM 

for  their  protection.  Apparently  other  physiological  nerve  pathways  exist, 
but  it  requires  a  stronger  sensory  stimulus  to  arouse  nerve  impulses  along 
these  paths.  In  explanation  we  may  suppose  that  the  stronger  afferent 
impulses  are  sufficient  to  overcome  the  resistance  of  increasingly  complex 
paths,  that  they  diffuse  through  greater  and  greater  extents  of  the  cord. 
But  we  may  repeat  that  in  the  normal  state  of  the  cord  this  diffusion  is  in  an 
orderly  physiological  sequence. 

Orderly  reflexes  can  be  called  forth  only  by  stimulating  sensory  nerve 
endings,  the  first  of  the  essential  structures  of  the  reflex  arc.  If  artificial 
stimuli  are  applied  to  a  nerve  trunk,  as  the  sciatic,  unco-ordinated  muscular 
responses  occur  because  the  sensory  stimuli  are  diffuse  and  general  and  are  not 
specific  and  local. 

An  involvement  of  multiple  pathways  may  also  be  accomplished  through 
decreasing  the  resistance  within  the  cord,  as  through  the  use  of  some  drug 
such  as  strychnine.  In  the  strychninized  frog  a  slight  stimulus  brings  about 
multiple  and  violent  reflex  spasms.  These  contractions  have  lost  their  order- 
liness and  are  unco-ordinated.  The  entire  musculature  contracts.  It  is  as 
though  the  strychnine  removed  all  differences  in  the  facility  with  which 
afferent  stimuli  spread  through  the  cord,  and  that  the  resistance  was  re- 
duced to  the  minimum.  The  strychnine  effect  is  possibly  due  to  a  decrease 
in  the  resistance  at  the  synapses,  and  possibly  also  to  an  increase  in  the 
irritability  of  the  discharging  nerve  cells. 

We  must  also  suppose  that  the  centers  are  particularly  sensitive  to  certain 
kinds  of  stimuli,  sometimes  producing  very  extensive  and  violent  muscular 
actions  in  response  to  a  slight  stimulus  of  a  special  kind.  Such  a  condition 
is  illustrated  in  the  violent  and  general  muscular  spasms  occurring  when  a 
small  particle  of  food  passes  into  the  larynx,  violent  expiratory  spasms 
accompanied  by  contractions  of  other  muscles  taking  place. 

The  time  taken  in  a  reflex  action  for  the  eye  in  man  has  been  found  to  be 
0.066  to  0.058  of  a  second,  but  this  estimate  includes  the  entire  time  from 
the  instant  of  stimulation  to  the  beginning  of  the  contraction  of  the  muscle. 

Functions  of  the  Spinal  Nerve  Roots. — The  anterior  spinal  nerve 
roots  are  efferent  in  function  and  the  posterior  are  afferent.  The  fact  is 
proved  in  various  ways.  Division  of  the  anterior  roots  of  one  or  more  nerves 
is  followed  by  complete  loss  of  motion  in  the  parts  supplied  by  the  fibers  of 
such  roots,  but  the  sensation  of  the  parts  remains  perfect.  Division  of  the 
posterior  roots  destroys  the  sensibility  of  the  parts  supplied  by  their  fibers, 
while  the  power  of  motion  continues  unimpaired.  Moreover,  stimulation  of 
the  ends  of  the  distal  portions  of  the  divided  anterior  roots  of  a  nerve  excites 
muscular  movements.  There  are  sometimes  slight  evidences  of  sensory  im- 
pulses due  to  recurrent  fibers  that  are  distributed  through  the  anterior  root 
to  the  spinal  meninges.  Stimulation  of  the  proximal  ends  of  the  anterior 
roots,which  are  still  in  connection  with  the  cord,  is  followed  by  no  appreciable 


SPINAL    REFLEXES    IN    MAN    AND    MAMMALS  595 

effect.  It  must  be  remembered,  however,  that  in  the  anterior  or  efferent 
nerves  other  fibers  besides  motor  are  contained,  e.g.,  vaso- motor,  secretory, 
heat  fibers,  and  when  the  distal  end  of  a  divided  nerve  is  stimulated,  the 
effects  are  exercised  not  only  upon  muscles,  but  upon  glands,  blood  vessels, 
etc.  Stimulation  of  the  distal  portions  of  the  divided  posterior  roots,  on  the 
other  hand,  produces  no  muscular  movements  and  no  manifestations  of 
pain;  for,  as  already  stated,  sensory  nerves  convey  impressions  only  toward 
the  nerve  centers.  Stimulation  of  the  proximal  portions  of  these  roots  elicits 
signs  of  intense  pain.  Muscular  movements  also  ensue;  but  these  are  the 
result  of  the  reflex  stimulation  of  the  motor  neurones  of  the  anterior  horn 
of  the  cord  or  are  movements  in  response  to  the  afferent  impulses  passing 
to  higher  centers  from  the  roots  stimulated. 

Functions  of  the  Ganglia  on  Posterior  Roots. — The  cells  of  the  posterior 
ganglia  act  as  centers  for  the  nutrition  of  the  nerve  fibers  given  off  from  them. 
When  these  are  cut,  the  parts  of  the  nerves  so  severed  degenerate,  while  the 
parts  which  remain  in  connection  with  the  cells  of  the  ganglia  do  not.  Thus 
on  section  of  the  posterior  nerve  root  beyond  the  ganglia  the  peripheral  part 
degenerates  and  the  central  does  not,  and  on  section  of  the  root  between 
the  ganglion  and  the  cord  the  central  part  degenerates  and  the  peripheral  is 
unaffected.  The  number  of  nerve  cells  in  the  spinal  ganglia  far  exceeds 
the  number  of  nerve  fibers  in  the  corresponding  root  (Hardesty).  The  extra 
cells,  see  figure  417,  serve  at  least  in  part  as  association  neurones  connecting 
the  bipolar  cells  of  the  ganglion,  cells  of  Dogiel,  and  to  connect  the  ganglion 
with  the  sympathetic  system  of  nerves. 

Spinal  Reflexes  in  Man  and  Mammals. — Much  of  our  knowledge  of 
the  reflexes  of  the  cord  is  derived  from  experiments  on  dogs,  though  paral- 
ysis of  the  lower  extremities  in  man,  by  accident  or  otherwise,  has  given  con- 
firmatory information.  In  man  the  spinal  cord  is  so  much  under  the  control 
of  the  higher  nerve  centers  that  its  own  individual  functions  in  relation  to  re- 
flex action  are  apt  to  be  overlooked.  But  if  the  skin  of  the  foot  is  stimulated, 
in  a  man  whose  lumbar  cord  is  completely  separated  by  injury  or  disease,  the 
foot  will  be  drawn  away  from  the  stimulus;  or,  if  the  stimulus  be  strong 
enough,  tne  entire  leg  will  be  moved.  In  both  cases  the  movement  may  be 
orderly  and  well  co-ordinated,  and  shows  that  the  sensory  stimulus  has  pro- 
duced a  co-ordinated  reflex  through  the  lumbar  cord.  The  injured  person 
feels  no  sensation  of  pain  nor  of  action,  and  the  phenomenon  is  independent 
of  the  higher  nerve  regions.  The  stimulus  that  is  supplied  to  man  must  be 
carefully  graded,  since  when  too  intense  it  calls  forth  muscular  spasms  or 
convulsive  action. 

When  the  cord  of  mammals  is  first  cut,  the  shock  is  very  great,  and  the 
lower  or  isolated  portion  of  the  cord  remains  for  a  time  quite  non-irritable. 
The  vaso- motor  and  thermogenic  centers  are  cut  off  from  the  periphery  so 
that  there  is  great  vascular  dilatation  and  marked  fall  of  temperature,  the 


596 


THE    NERVOUS    SYSTEM 


effects  of  which  are  likely  to  lead  to  death  unless  the  operated  animal  is 
carefully  attended.  But  these  effects  are  slowly  recovered  from,  and  man, 
as  well  as  the  lower  mammals,  soon  regains  the  vascular  tone.  The  general 
tonus  of  the  muscular  system,  which  is  lost  at  first,  is  also  regained. 

In  this  partially  recovered  condition  man,  and  such  animals  as  the  cat,  the 
dog,  and  the  monkey,  perform  certain  of  the  lower  functions  with  a  re- 
markable degree  of  perfection.  Of  course  these  functions  are  under  constant 
co-ordinative  regulation  and  control  in  the  normal  animal,  but  experiments 


FIG.  370. — Scheme  of  the  Relation  of  the  Posterior  Root  Fibers  upon  Entering  the 
Cord.  A,  The  branch  of  the  dorsal  root  fibers  upon  entering  cord;  B,  terminal  arborization 
about  cell  bodies  of  the  cord;  DR,  axones  of  the  dorsal  root;  B,  their  ascending  and  descend- 
ing branches;  C,  collaterals.  (After  Cajal.) 

and  observation  have  shown  that  much  of  such  activity  really  is  a  primary 
function  of  the  cord.  Of  these  activities  the  following  may  be  especially 
mentioned:  Muscular  tonus,  general  reflexes,  the  special  reflexes  of  mic- 
turition, defecation,  erection  and  the  sexual  reflex,  and  parturition,  some 
of  which  will  be  briefly  discussed. 

The  Center  of  the  Tone  of  Muscles. — The  tonic  influence  of  the  spinal  cord 
on  the  sphincter  ani  and  sphincter  urethras  will  be  presently  mentioned.  The 
cord  maintains  these  muscles  in  permanent  tonic  contraction.  The  condition 
of  the  sphincters,  however,  is  not  altogether  exceptional.  Their  contraction 


SPINAL   REFLEXES    IN   MAN   AND    MAMMALS  597 

is  the  same  in  kind,  though  it  exceeds  in  degree,  that  condition  of  the  voluntary 
muscles  which  has  been  called  tone,  a  condition  of  slight  contraction  which 
they  always  maintain  during  health.  This  tone  of  all  the  muscles  of  the 
trunk  and  limbs  depends  on  the  spinal  cord,  just  as  does  the  contraction  of 
the  sphincters.  If  an  animal  be  killed  by  injury  or  removal  of  the  brain, 
the  muscles  retain  their  tenseness,  but  if  the  spinal  cord  be  destroyed,  all  the 
muscles  become  loose,  flabby,  and  atonic,  remaining. so  till  rigor  mortis 
commences. 

If  an  animal,  such  as  the  dog,  be  held  off  the  ground  in  the  erect  position 
assumed  by  the  human  body,  when  the  trunk  and  hind  limb  muscles  are  not 
in  voluntary  contraction  the  limbs  will  assume  a  normal  pendular  position. 
In  the  pendular  position  the  legs  of  a  dog  with  cord  severed  hang  more  limp 
and  are  more  completely  extended.  The  muscles  of  the  former  exhibit  that 
tone  which  keeps  antagonistic  muscles  always  slightly  tense,  the  muscles  of 
the  latter  have  lost  the  tenseness. 

Whether  or  not  muscular  tone  is  maintained  through  the  constant  sub- 
minimal  action  of  sensory  nerve  impulses  on  the  tonic  centers  of  the  cord,  or 
whether  these  centers  are  automatic  in  their  action,  is  a  question  that  can  be 
answered  only  by  inference.  The  probability  is  that  tone  is  a  reflex  activity, 
though  it  may  be  contributed  to  by  the  normal  healthy  nutritional  condition 
of  the  muscles  themselves — a  condition  which  is  itself  dependent  on  the 
trophic  influence  of  the  nerve  cells  of  the  cord  and  brain  stem. 

The  Ano-spinal  or  Defecation  Center. — The  mode  of  action  of  the  ano- 
spinal  center  appears  to  be  this:  The  mucous  membrane  of  the  rectum  is 
stimulated  by  the  presence  of  feces  or  of  gas  in  the  large  bowel.  The  stim- 
ulus passes  up  by  the  afferent  nerves  of  the  hemorrhoidal  and  inferior  mesen- 
teric  plexuses  to  the  center  situated  in  the  lumbar  enlargement  of  the  cord, 
and  is  reflected  through  the  pudendal  plexus  to  the  anal  sphincter,  and  to  the 
muscular  tissue  in  the  wall  of  the  lower  bowel.  In  this  way  there  is  produced 
a  relaxation  of  the  first  and  a  contraction  of  the  second,  and  expulsion  of  the 
contents  of  the  bowel  follows.  The  center  in  the  spinal  cord  is  partially 
under  the  control  of  the  will,  so  that  its  action  may  be  either  inhibited  or 
augmented.  The  action  may  be  helped  by  the  abdominal  muscles,  which 
are  voluntary  muscles,  but  are  also  stimulated  to  contract  by  reflex  action. 

The  Vesico-spinal  or  Micturition  Center. — The  vesico-spinal  center  acts 
in  a  very  similar  way  to  that  of  the  ano-spinal.  The  center  is  also  in  the 
lumbar  enlargement  of  the  cord.  It  is  stimulated  to  action  reflexly  by  the 
presence  of  urine  in  the  bladder.  The  action  may  be  voluntary  and  is  excited 
by  the  sensation  of  distention  of  the  bladder  by  the  urine.  The  sensory  fibers 
concerned  are  the  posterior  roots  of  the  lower  sacral  nerves.  The  action  of 
the  spinal  center  is  double,  or  it  may  be  supposed  that  the  center  consists  of 
two  parts,  one  of  which  is  usually  in  action  and  maintains  the  tone  of  the 
sphincter,  and  the  other  which  causes  contraction  of  the  bladder  and  other 


THE    NERVOUS    SYSTEM 

muscles.  When  evacuation  of  the  bladder  occurs  sensory  impulses  pass  to 
that  part  of  the  center  which  discharges  impulses  to  the  bladder  and  to  cer- 
tain accessory  muscles  which  cause  their  contraction;  and  impulses  pass  to 
that  part  of  the  center  which  inhibits  the  tonic  action  on  the  sphincter  ure- 
thrae  which  procures  its  relaxation.  The  way  having  been  opened  by  the 
relaxation  of  the  sphincter,  the  urine  is  expelled  by  the  combined  action  of 
the  bladder  and  accessory  muscles.  The  cerebrum  may  exert  its  influence 
on  the  reflex  not  only  by  stimulating  the  center  to  action,  but  also  by  inhibit- 
ing its  action. 

The  Genito-spinal  Center. — The  presence  of  the  genito-spinal  center  is 
proven  by  the  fact  that  dogs,  and  even  man,  are  known  to  discharge  semen 
when  the  lumbar  cord  is  severed  and  all  voluntary  motion  and  sensibility  are 
lost.  The  center  situated  in  the  lumbar  enlargement  of  the  spinal  cord  is 
stimulated  to  action  by  sensory  impressions  from  the  glans  penis.  Efferent 
impulses  from  the  center  excite  the  successive  and  co-ordinate  contractions  of 
the  muscular  fibers  of  the  vasa  deferentia  and  vesiculae  seminales  and  of  the 
accelerator  urinae  and  other  muscles  of  the  urethra;  and  a  forcible  expulsion 
of  semen  takes  place,  which,  in  cases  of  paraplegia,  are  not  felt. 

The  Erected  Center. — This  center  is  also  situated  in  the  lumbar  region  and 
is  a  vascular  center,  already  described  in  the  chapter  on  Circulation.  It 
is  reflexly  excited  to  action  by  the  sensory  nerves  of  the  penis,  and  also  in  the 
normal  animal  by  impulses  passing  down  from  the  cerebrum.  Efferent 
impulses  produce  dilatation  of  the  vessels  of  the  penis. 

The  Parturition  Center. — The  center  for  the  expulsion  of  the  contents  of 
the  uterus  in  parturition  is  situated  in  the  lumbar  spinal  cord  rather  higher 
up  than  the  other  centers  already  enumerated.  The  stimulation  of  the 
uterus  may,  under  certain  conditions,  excite  the  center  to  send  out  impulses 
which  produce  a  contraction  of  the  uterine  walls  and  expulsion  of  the  con- 
tents of  the  cavity.  The  center  is  independent  of  the  will  since  delivery 
takes  place  in  paraplegic  women,  and  also  while  a  patient  is  under  the  influ- 
ence of  chloroform.  Again,  as  in  the  cases  of  defecation  and  micturition,  the 
abdominal  and  thoracic  muscles  assist;  their  action  being  for  the  most  part 
reflex  and  involuntary. 

Inhibition  of  Reflex  Actions. — Movements  such  as  are  produced  by  stimu- 
lating the  skin  of  the  lower  extremities  in  the  human  subject,  after  division 
or  disorganization  of  a  part  of  the  spinal  cord,  do  not  always  occur  when  the 
cerebrum  is  active  and  the  connection  between  the  cord  and  the  brain  is  in- 
tact. The  reflex  which  would  occur  in  the  animal  with  spinal  cord  only  is 
suppressed  or  inhibited  in  the  normal  animal  through  the  regulative  action  of 
the  higher  cerebral  centers.  When  one  is  anxiously  thinking,  even  slight 
stimuli  may  produce  involuntary  and  reflex  movements.  So,  also,  during 
sleep,  such  reflex  movements  may  be  observed,  when  the  skin  is  touched  or 
tickled;  for  example,  when  one  touches  the  palm  of  the  hand  of  a  sleeping 


CUTANEOUS    AND    MUSCLE    REFLEXES    AS    DIAGNOSTIC    SIGNS        599 

child,  the  impression  on  the  skin  of  the  palm  producing  a  reflex  movement  of 
the  muscles  which  close  the  hand.  But  when  the  individual  is  awake  no 
such  reflex  is  produced. 

Further,  many  reflex  actions  are  capable  of  being  more  or  less  controlled 
or  even  altogether  prevented  by  the  will,  of  which  the  following  may  be 
quoted  as  familiar  examples: 

When  the  foot  is  tickled  we  can,  by  an  effort  of  will,  prevent  the  reflex 
action  of  jerking  it  away.  So,  too,  the  involuntary  closing  of  the  eyes  and 
starting  back,  when  a  blow  is  aimed  at  the  head,  can  be  similarly  restrained. 
Darwin  has  mentioned  an  interesting  example  of  the  way  in  which  such  an 
instinctive  reflex  act  may  override  the  strongest  effort  of  the  will.  He  placed 
his  face  close  against  the  glass  of  the  cobra's  cage  in  the  Reptile  House  at 
the  Zoological  Gardens,  and,  though  of  course  thoroughly  convinced  of  his 
perfect  security,  could  not  by  any  effort  of  the  will  prevent  himself  from 
starting  back  when  the  snake  struck  with  fury  at  the  glass. 

It  can  be  readily  shown,  by  comparing  a  spinal  frog  and  a  normal  unin- 
jured frog,  that  stimuli  which  call  forth  definite  reflexes  in  the  first  often 
produce  no  movement  of  the  second. 

Cutaneous  and  Muscle  Reflexes  as  Diagnostic  Signs. — In  the  hu- 
man subject  two  classes  of  reflex  actions  dependent  upon  the  spinal  cord  are 
usually  distinguished,  the  alterations  of  which,  either  of  increase  or  of  diminu- 
tion, are  indications  of  some  abnormality,  and  are  used  as  a  means  of  diag- 
nosis in  nervous  and  other  disorders.  They  are  termed,  respectively,  cutane- 
ous reflexes  and  muscle  reflexes.  Cutaneous  reflexes  are  set  up  by  a  gentle 
stimulus  applied  to  the  skin.  The  subjacent  muscle  or  muscles  contract  in 
response.  Although  these  cutaneous  reflex  actions  may  be  demonstrated 
almost  anywhere,  yet  certain  of  such  actions  as  being  most  characteristic  are 
distinguished,  e.g.,  plantar  reflex,  gluteal  reflex,  i.e.,  a  contraction  of  the 
gluteus  maximus  when  the  skin  over  it  is  stimulated;  cremaster  reflex,  re- 
traction of  the  testicle  when  the  skin  of  the  inside  of  the  thigh  is  stimulated, 
and  the  like.  The  ocular  reflexes,  too,  are  important.  They  are  contraction 
of  the  iris  on  exposure  to  light,  and  its  dilatation  on  stimulating  the  skin  of 
the  cervical  region.  All  of  these  cutaneous  reflexes  are  true  reflex  actions. 
They  differ  in  different  individuals,  and  are  more  easily  elicited  in  the  young. 

Muscle  reflexes  or  tendon  reflexes  consist  of  a  contraction  of  a  muscle 
under  conditions  of  more  or  less  tension,  when  its  tendon  is  sharply  tapped. 
The  so-called  patellar-tendon  reflex  " knee-jerk"  is  the  best  known  of  this 
variety  of  reflexes.  If  one  knee  be  slightly  flexed,  as  by  crossing  it  over  the 
other,  so  that  the  quadriceps  femoris  is  extended  to  a  moderate  degree,  and 
the  tendon  of  the  patella  be  tapped  with  the  fingers,  the  muscle  contracts  and 
the  foot  is  jerked  forward.  Another  variety  of  the  same  phenomenon  is  seen 
if  the  foot  is  flexed  so  as  to  stretch  the  calf  muscles,  and  the  tendo  Achillis  is 
tapped;  the  foot  is  extended  by  the  contraction  of  the  stretched  muscles.  It 


6oo 


THE    NERVOUS    SYSTEM 


appears,  however,  that  the  tendon  reflexes  are  not  exactly  what  their  name 
implies.  The  interval  between  the  tap  and  the  contraction  is  said  to  be  too 
short  for  the  production  of  a  true  reflex  action.  It  is  suggested  that  the  con- 
traction is  caused  by  local  stimulation  of  the  muscle,  but  that  this  would  not 
occur  unless  the  muscle  had  previously  been  stimulated  by  the  tension  applied, 
and  placed  in  a  condition  of  excessive  irritability.  It  is  probable  that  the 

condition  on  which  it  depends  is  a  reflex  change 
in  the  spinal  irritability  acting  on  the  muscle  or 
exaggerated  muscular  tone,  which  is  admitted 
to  be  a  reflex  phenomenon  in  the  spinal  cord. 
Conduction  in  the  Spinal  Cord. — With 
the  differentiation  of  the  central  nerve  axis  in 
vertebrates  the  conduction  in  the  spinal  cord 
becomes  of  increasing  importance,  reaching  its 
maximum  in  man.  It  is  evident  that  the 
cord  is  the  path  by  which  all  nerve  impulses 
arising  in  the  trunk  or  in  the  arms  and  legs 
must  reach  the  brain,  or  vice  versa.  Impulses 
of  peripheral  origin  can  and  do  produce  re- 
flexes, but  they  can  arouse  sensations  and  be 
perceived  only  after  they  have  been  conducted 
to  the  cerebral  cortex.  Motor  impulses  arising 
in  the  brain  can  reach  the  motor  cells  of  the 
anterior  columns  of  the  cord  only  through  the 
cord  as  a  conducting  path.  The  continuity  of 
the  cord,  therefore,  while  not  necessary  for  the 
execution  of  reflexes,  is  absolutely  necessary 
for  the  higher  co-ordinations  of  the  reflexes 
and  for  the  excitation  and  controlling  influence 
of  the  brain. 

Illustrations  of  this  are  furnished  by  various 

Columns  of  the  Cord.  (Edinger.)    examples  of  paralysis,  but  by  none  better  than 

by  the  common  paraplegia,  or  loss  of  sensation 

and  voluntary  motion  in  the  lower  part  of  the  body,  in  consequence  of 
destructive  disease  or  injury  of  a  section  including  the  whole  thickness  of 
the  spinal  cord.  Such  lesions  destroy  the  communication  between  the  brain 
and  all  parts  of  the  spinal  cord  below  the  seat  of  injury,  and  consequently 
cut  off  from  their  connection  with  the  brain  the  various  organs  supplied 
with  nerves  issuing  from  those  parts  of  the  cord. 

It  is  not  probable  that  the  conduction  of  motor  and  of  sensory  impulses  is 
effected  under  ordinary  circumstances,  to  so  great  an  extent  as  was  formerly 
supposed,  through  the  gray  substance  of  the  cord,  i.e.,  from  cell  to  cell 
through  the  short  filaments  lying  wholly  within  the  gray  substance.  But 


FIG.  371. — Diagram  to  Show 
the  Manner  in  which  the  Fibers 
of  the  Posterior  Nerve  Roots 
Enter  and  Ascend  the  Posterior 


SENSORY   IMPULSES  6oi 

cells  with  fibers  running  for  short  distances  in  the  ground  bundles  are 
numerous,  and  these  short  connectives  are  capable  of  conducting  impulses 
along  the  cord.  All  parts  of  the  cord  are  not  alike  able  to  conduct  all  im- 
pressions; and  as  there  are  separate  nerve  fibers  for  motor  and  for  sensory 
impressions,  so  in  the  cord  separate  and  determinate  tracts  serve  to  conduct 
always  the  same  kind  of  impressions.  The  sensations  of  touch,  and  perhaps 
of  temperature  and  pain,  do  not  appear  to  have  such  sharply  limited  tracts 
as  do  the  motor  impulses. 

Experimental  and  other  observations  point  to  the  following  conclusions 
regarding  the  conduction  of  sensory  and  motor  impressions  through  the 
spinal  cord.  Many  of  these  conclusions  must,  however,  be  received  with 
considerable  reserve. 

Sensory  Impulses. — The  sensory  impressions  of  couch,  pain,  heat  and 
cold,  and  of  the  muscle  sense  are  conducted  to  the  spinal  cord  by  the  posterior 
nerve  roots.  Certain  sensory  impressions  are  then  carried  directly  into  the 
fasciculus  gracilis  on  the  same  side,  and  thence  up  to  the  nucleus  of  this 
column  in  the  medulla.  It  is  mainly  the  impulses  of  the  muscle  sense  and 
of  the  sense  of  touch  that  take  this  course  through  the  cord,  though  the 
sense  of  touch  is  not  wholly  interrupted  upon  injury  to  the  posterior  columns. 
In  lower  animals  it  is  scarcely  interfered  with  at  all.  The  posterior  funiculi 
unquestionably  are  the  primary  muscle  sensory  paths.  Visceral  sensations 
are  carried  by  the  posterior  root  fibers  to  the  cells  of  the  column  of  Clarke 
in  the  posterior  horn,  figure  363.  From  there  the  impulses  pass  to  the  direct 
cerebellar  tract  on  the  same  side,  and  thence  up  through  the  medulla  to  the 
cerebellum,  figure  388.  The  impressions  of  pain,  and  of  heat  and  cold,  are 
conveyed  to  the  nerve  cells  in  the  posterior  cornua  of  the  same  side  in  part, 
and  in  part  to  the  nerve  cells  in  the  posterior  cornu  and  median  gray  matter 
of  the  apposite  side.  From  this  point,  the  impulses  are  taken  up  again  by 
intermediary  neurones  and  conveyed  through  the  anterior  and  lateral  columns 
of  the  cord  to  the  brain  in  the  ascending  superficial  antero-lateral  tract,  or 
tract  of  Gowers.  By  reason  of  the  great  number  of  collaterals  and  the 
interpolation  in  the  course  of  the  sensory  path  of  many  intermediary  neu- 
rones, it  has  been  difficult  to  make  out  very  sharply  defined  tracts  in  the 
spinal  cord  for  the  conduction  of  the  sensations  of  temperature,  pain,  and 
touch.  If  one  set  of  fibers  is  destroyed  by  disease,  others  seem  able,  through 
the  collaterals,  to  take  up  its  functions.  We  can  say  that  injury  to  the 
lateral  columns  has  resulted  in  loss  of  the  sense  of  pain,  heat,  and  cold,  but 
with  only  partial  disturbance  of  touch  sensations. 

It  is  probable,  also,  that  pain  and  temperature  sensations  cross  over  at 
once  to  a  considerable  extent  and  pass  up  in  the  opposite  side  of  the  cord  to 
which  they  enter.  Touch  and  the  muscle  sense  impressions,  especially  the 
latter,  pass  up  largely  upon  the  same  side  until  they  reach  the  medulla  or 
cerebellum. 


602  THE    NERVOUS    SYSTEM 

Motor  Impulses. — Motor  impulses  are  conveyed  downward  from  the 
cerebral  cortex  of  the  brain  along  the  pyramidal  tracts,  viz.,  the  lateral 
and  the  anterior,  chiefly  the  former.  In  the  lateral  pyramidal  tract  the 
impressions  pass  down  chiefly  on  the  side  opposite  to  which  they  originate, 
having  crossed  over  in  the  decussation  in  the  medulla.  But  some  motor 
impulses  do  not  cross  in  the  medulla,  but  descend  in  the  anterior  pyramidal 
tract  to  lower  levels  of  the  cord,  where  they  cross  in  the  anterior  commissure. 
The  motor  fibers  for  the  legs  partially  pass  downward  in  the  lateral  columns 
of  the  same  side  without  decussation.  This  is  also  probably  the  case  with 
the  bilateral  muscles,  i.e.,  muscles  of  the  two  sides  that  act  together,  such 
as  the  intercostal  muscles  and  other  muscles  of  the  trunk. 

It  is  quite  certain,  as  was  just  now  pointed  out,  that  the  fibers  of  the  an- 
terior nerve  roots  are  more  numerous  than  the  fibers  proceeding  downward 
from  the  brain  in  the  pyramidal  tracts,  the  so-called  pyramidal  fibers.  This 
is  because  each  pyramidal  fiber  is  really  a  very  long  nerve  process  or  axone, 
and  is  supplied  in  its  course  with  a  large  number  of  collaterals.  These  go  off 
at  different  points,  and  thus  put  it  in  relation  with  different  groups  of  nerve 
cells  in  the  anterior  columns  at  various  levels.  Each  nerve  fiber  of  the 
pyramidal  tract,  by  means  of  its  collaterals,  can  control  a  number  of  nerve 
cells,  and  can  thus  co-ordinate  the  action  of  impulses  sent  out  through  the 
anterior  roots  to  a  number  of  groups  of  muscles.  In  other  words,  the  gray 
matter  of  the  anterior  columns  contains  an  apparatus  with  various  com- 
plicated co-ordinating  powers,  which  apparatus  is  under  the  regulative 
control  of  the  neurones  whose  cells  of  origin  are  in  the  cortex  of  the  cerebrum. 
This  is  the  same  apparatus  that  is  also  reflexly  influenced  by  sensory  im- 
pressions passing  to  the  cord  from  the  periphery. 

Division  of  a  single  anterior  pyramid  of  the  medulla  at  a  point  just  above 
the  decussation  is  followed  by  paralysis  of  voluntary  motions  in  the  muscles  of 
the  opposite  side  in  all  parts  below.  Disease  or  division  of  any  part  of  the 
cerebro-spinal  axis  below  the  seat  of  decussation  of  the  pyramids  is  followed 
by  impairment  or  loss  of  voluntary  motion  on  the  same  side  of  the  body. 
The  paralysis  is  never  quite  complete,  and  the  opposite  side  usually  shows 
some  slight  impairment  of  function  of  the  muscle. 

When  one-half  of  the  spinal  cord  is  cut  through  in  monkeys,  the  results 
are  as  follows  (Mott):  Motor  paralysis  of  the  muscles  of  the  same  side 
(never  complete  paralysis  of  the  muscles  used  in  bilateral  associated  action), 
followed  by  gradual  recovery  of  muscular  movement,  except  of  the  finer 
movements  of  the  hand  and  foot;  wasting  and  flabbiness  of  the  muscles; 
partial  sensory  paralysis  of  the  same  side  (temperature,  touch,  pain,  and 
pressure) ;  temporary  vaso-motor  paralysis  on  the  same  side.  The  tempera- 
ture of  the  affected  side  is  depressed  i  to  3°  F. 


THE   BRAIN 


603 


THE  BRAIN. 

General  Arrangement  of  Parts  of  the  Brain. — The  great  relative  and 
absolute  size  of  the  cerebral  hemispheres  in  the  adult  man  and  in  mammals 
to  a  great  extent  mask  the  real  arrangement  of  the  several  parts  of  the  brain. 
An  examination  of  the  accompanying  diagram,  figures  370,  371,  reveals  that 
the  parts  of  the  brain  are  disposed  in  a  linear  series,  as  follows  (from  before 
backward):  Olfactory  lobes,  cerebral  hemispheres,  thalamencephalon 
(thalami  and  third  ventricle),  the  mid-brain  (corpora  quadrigemina  and 
crura  cerebri,  medulla  oblongata,  and  cerebellum. 


FIG.  372. — Diagrammatic  Horizontal  Section  of  the  Vertebrate  Brain.  The  figures 
serve  both  for  this  and  the  next  diagram.  Mb,  mid-brain;  what  lies  in  front  of  this  is  the 
fore-,  and  what  lies  behind  the  hind-brain;  Lt,  lamina  terminalis;  Olf,  olfactory  lobes; 
Hemp,  hemispheres;  Th.  E,  thalamencephalon;  Pn,  pineal  gland;  Pyt  pituitary  body; 
F.  M.,  foramen  of  Munro;  cs,  corpus  striatum;  Th,  optic  thalamus,  CC\  crura  cerebri;  the 
mass  lying  above  the  canal  represents  the  corpora  quadrigemina;  Cb,  cerebellum;  I-IX,  the 
nine  pairs  of  cranial  nerves;  i,  olfactory  ventricle;  2,  lateral  ventricle;  3,  third  ventricle;  4, 
fourth  ventricle;  + ,  iter  a  tertio  ad  quartum  ventriculum.  (Huxley.) 


The  linear  arrangement  of  parts  actually  occurs  in  an  early  stage  of  the 
development  of  the  human  fetus,  and  it  is  permanent  in  some  of  the  lower 
vertebrata.  In  fishes  the  cerebral  hemispheres  are  represented  by  a  pair 
of  ganglia  intervening  between  the  olfactory  and  the  optic  lobes,  and  con- 
siderably smaller  than  the  latter,  their  adult  development  is  fairly  well  repre- 
sented by  the  figure  373.  In  Amphibia  the  cerebral  lobes  are  further 
developed,  and  are  larger  than  any  of  the  other  ganglia. 


604 


THE    NERVOUS    SYSTEM 


In  reptiles  and  birds  the  cerebral  ganglia  attain  a  still  further  development 
and  in  Mammalia  the  cerebral  hemispheres  exceed  in  weight  all  the  rest  of  the 
brain.  As  we  ascend  the  scale,  the  relative  size  of  the  cerebrum  increases,  till 


EL  v 


FlG.  373. — Longitudinal  and  Vertical  Diagrammatic  Section  of  a  Vertebrate  Brain. 
Letters  as  before.  Lamina  terminalis  is  represented  by  the  strong  black  line  joining  Pn 
and  Py.  (Huxley.) 


FIG.  374. — Base  of  the  Brain,  i,  Superior  longitudinal  fissure;  2,  2',  2",  anterior 
cerebral  lobe;  3,  fissure  of  Sylvius,  between  anterior  and  4,  4',  4",  middle  cerebral  lobe; 
5»  5'»  posterior  lobe;  6,  medulla  oblongata.  The  figure  is  in  the  right  anterior  pyramid;  7, 
8,  9,  10,  the  cerebellum;  +,  the  inferior  vermiform  process.  The  figures  from  7.  to  IX. 
are  placed  against  the  corresponding  cerebral  nerves;  777.  is  placed  on  the  right  cms  cerebri. 
VI.  and  F77.  on  the  pons  Varolii;  X.,  the  first  cervical  or  suboccipital  nerve. 


Thomson.)     X 


(Allen 


in  the  higher  apes  and  man  the  hemispheres,  which  commenced  as  two  little 
lateral  buds  from  the  anterior  cerebral  vesicle,  having  grown  upward  and 
backward,  completely  covering  in  and  hiding  from  view  practically  all  the 


THE  BRAIN    OR   ENCEPHALON  605 

rest  of  the  brain.  At  the  same  time  the  smooth  surface  of  the  cerebral 
cortex  of  many  lower  mammalia,  such  as  the  rabbit,  is  replaced  by  the  laby- 
rinth of  convolutions  of  the  human  brain. 

When  the  cerebral  hemispheres  are  removed,  several  large  basal  masses 
of  nerve  substance  are  revealed:  the  optic  thalami,  the  corpora  quadrigemina, 
and  the  cms  cerebri.  These  structures,  together  with  the  pons  and  the  me- 
dulla, form  a  direct  continuation  forward  of  the  spinal  cord  and  sometimes 
are  designated  under  the  general  term  of  the  brain  stem. 


FIG.  375. — Plan  in  Outline  of  the  Brain  as  seen  from  the  Right  Side.  X  $.  The  parts 
are  represented  as  separated  from  one  another  somewhat  more  than  natural,  so  as  to  show 
their  connections.  A,  Cerebrum;  /,  g,  h,  its  anterior,  middle,  and  posterior  lobes  e, 
fissure  of  Sylvius;  B,  cerebellum;  C,  pons  Varolii;  D,  medulla  oblongata;  a,  peduncles  of  the 
cerebrum;  6,  c,  d,  superior,  middle,  and  inferior  peduncles  of  the  cerebellum.  (From 
Quain.) 


The  human  brain  on  superficial  examination  does  not  seem  to  follow  the 
general  plan  outlined  above,  but  when  the  cerebral  hemispheres  and  the 
cerebellum  are  removed  then  it  is  found  that  what  remains  closely  follows 
the  plan  presented.  This  central  axis  is  shown  in  part  in  figure  379. 

The  morphological  parts  of  the  brain  usually  given  are: 

The  Brain  or  Encephalon. 

I.  The  hind-brain  or  rhombencephalon. 

1.  Myelencephalon. 

a.  Bulb  or  medulla  oblongata. 

2.  Metencephalon. 

b.  Pons  Varolii. 

c.  Cerebellum. 


606  THE   NERVOUS    SYSTEM 

II.  Mid-brain  or  mesencephalon. 

d.  Corpora  quadrigemina 

e.  Crura  cerebri. 

III.  Fore-brain,  prosencephalon. 

3.  Thalamencephalon. 

f.  Optic  thalami. 

4.  Telencephalon. 

g.  Corpora  geniculata. 
h.  Corpora  striata. 

i.  Cerebral  hemispheres. 

THE  MEDULLA  OBLONGATA  AND  PONS. 
STRUCTURE. 

Anatomical  Structure. — The  medulla  oblongata  is  continuous  with 
the  spinal  cord  at  the  upper  end.  It  lies  within  the  cranial  cavity  and  forms 
the  first  part  of  the  brain  stem.  The  medulla  consists  of  masses  of  nerve 
cells  situated  in  the  interior,  but  pretty  generally  distributed  throughout 
the  mass.  The  cell-masses  are  subdivided  by  laminae  of  nerve  fibers  into 
groups,  or  nuclei,  which  give  origin  to  or  form  the  terminations  of  the  various 
ranks  of  nerve  fibers. 

The  nerve  fibers  are  arranged  partly  in  columns  and  partly  in  fasciculi 
traversing  the  central  cellular  matter.  The  medulla  oblongata  is  larger  than 
any  part  of  the  spinal  cord.  Its  columns  are  pyriform,  enlarging  as  they  pro- 
ceed toward  the  upper  part  of  the  brain,  and  are  continuous  with  funiculi 
of  the  spinal  cord.  Each  half  of  the  medulla,  therefore,  may  be  divided 
into  three  columns  or  tracts  of  fibers,  continuous  with  the  three  columns  of 
funiculi  or  of  the  spinal  cord.  The  columns  are  more  prominent  than  those 
of  the  spinal  cord,  and  are  separated  from  each  other  by  deeper  grooves. 
The  anterior,  continuous  with  the  anterior  columns  of  the  cord,  are  called 
the  pyramids.  The  postero-median  and  external  are  represented  at  the 
posterior  or  dorsal  aspect  of  the  cord  as  the  fasciculus  gracilis  and  the  fascic- 
ulus cuneatus.  The  posterior  pyramids  of  the  medulla,  which  include  these 
two  columns  of  white  matter,  soon  become  much  increased  in  width  by  the 
addition  of  a  new  column  of  white  matter  outside  the  other  two,  which  is 
known  as  the  fasciculus  of  Rolando.  In  the  upper  portion  of  the  medulla  the 
fasciculi  are  replaced  by  the  restiform  bodies  (the  inferior  peduncles  of  the 
cerebellum).  The  lateral  columns  of  the  cord  are  scarcely  represented  as 
such  in  the  bulb. 

It  may  be  said  then  that  the  bulb  at  its  commencement  differs  only  slightly 
in  size  from  the  cord,  with  which  it  is  continuous.  It  soon  becomes  larger 
both  laterally  and  antero-posteriorly.  It  opens  out  on  the  dorsal  surface  into 


ANATOMICAL    STRUCTURE 


607 


a  space  which  is  known  as  the  fourth  ventrical,  and  from  being  a  cylinder 
with  a  central  canal  it  is  flattened  out  on  the  dorsal  surface  by  the  gradual 
approach  of  the  central  canal  to  that  surface,  where  it  is  directly  continuous 
with  the  fourth  ventricle. 

On  the  anterior  or  ventral  surface  of  the  bulb  it  is  found  that  the  anterior 
fissure  is  occupied  at  the  most  posterior  part  by  fibers  which  are  crossing 
from  one  side  to  the  other.  This  is  known  as  the  decussation  of  the  pyramids. 
It  is  formed  of  the  lateral  pyramidal  fibers.  The  lateral  pyramidal  fibers  of 


FIG.  377. 

FIG.  376. — Ventral  or  Anterior  Surface  of  the  Pons  Varolii  and  Medulla  Oblongata, 
a,  a,  Anterior  pyramids;  b,  their  decussation;  c,  c,  olivary  bodies;  d,  d,  restiform  bodies;  e, 
arciform  fibers;/,  fibers  passing  from  the  anterior  column  of  the  cord  to  the  cerebellum;  g. 
anterior  column  of  the  spinal  cord;  h,  lateral  column;  p,  pons  Varolii;  i,  its  upper  fibers; 
5,  5,  roots  of  the  fifth  pair  of  nerves. 

FIG.  377. — Dorsal  or  Posterior  Surface  of  the  Pons  Varolii,  Corpora  Quadrigemina, 
and  Medulla  Oblongata.  The  peduncles  of  the  cerebellum  are  cut  short  at  the  side, 
a,  a,  the  upper  pair  of  corpora  quadrigemina;  b,  b,  the  lower;/,/,  superior  peduncles  of  the 
cerebellum;  c,  eminence  connected  with  the  nucleus  of  the  hypoglossal  nerve;  e,  that  of  the 
glosso-pharyngeal  nerve;  i,  that  of  the  vagus  nerve;  d,  d,  funiculus  cuneatus;  p,  p,  funiculus 
gracilis;  v,  v,  groove  in  the  middle  of  the  fourth  ventricle,  ending  below  in  the  calamus 
scriptorius;  7,  7,  roots  of  the  auditory  nerves. 

either  side  after  crossing  the  middle  line  become  part  of  the  pyramid  of  the 
opposite  side;  the  rest  of  the  pyramid  is  made  up  of  the  fibers  which  in  the 
anterior  column  of  the  cord  are  known  as  the  direct  or  anterior  pyramidal 
tract.  The  pyramidal  fibers  are  those  which  degenerate  after  lesions  of  the 
parts  of  the  cerebrum  known  as  the  motor  areas  of  the  cortex.  They  are 
the  descending  fibers  of  communication  between  the  cerebral  motor  cells 
of  the  cortex  and  the  different  segments  of  the  spinal  cord.  The  outer  bor- 
ders of  the  anterior  pyramids  of  the  bulb  are  marked  by  the  exit  from  that 
part  of  the  nervous  axis  of  the  twelfth  or  hypoglossal  nerve.  Still  more  later- 


6o8 


THE    NERVOUS    SYSTEM 


ally  there  is  on  either  side  a  rounded  elevation  or  column,  the  olivary  body. 
It  begins  at  a  level  a  little  lower  than  the  opening  of  the  fourth  ventricle.  On 
the  dorsal  side  of  the  olivary  body  is  the  line  of  origin  of  the  eleventh,  tenth, 
and  ninth  nerves,  and  from  this  to  the  posterior  fissure  is  the  region  corre- 
sponding to  the  lateral  and  posterior  columns  of  the  cord. 

The  changes  in  structure  which  are  noticed  in  a  series  of  sections  of  the 
bulb  from  below  upward  may  be  summarized:  In  the  dorsal  or  posterior  re- 
gion, the  posterior  columns  are  pushed  more  to  each  side  by  the  large  number 
of  sensory  fibers  ascending  in  the  posterior  funiculus  and  terminating  in  the 


Optic  chiasma 


Optic 

Corpus  geniculatum 
extern  um 

Corpus  geniculatum 

internum 

Locus  perforatus 

posticus 


Middle  peduncle 
of  tbe  cerebellum 


Restiform  body 

Oli 

Pyramid 

Anterior  superficia 
arcuate  fibres 

Decussation 
pyramids 


Optic  nerve 
Infundibulum 
Tuber  cinereum 
Corpora  mammillaria 
culo-motor  nerve  (III.) 

Trochlear  nerve  (IV.) 
winding  round  the  crus 
cerebri 

Trigeminal  nerve  (V.) 

Abducent  nerve  (VI.) 
Facial  nerve  (VII.) 
Auditory  nerve  (VIII.) 

Vago-glossopharyngeal 
nerve  (IX.  and  X.) 

Hypoglossal 
nerve  (XII.) 

.Spinal  accessory 
nerve  (XI.) 

First  cervical  nerve 


FIG.  378. — Front  View  of  the  Medulla,  Pons,  and  Mesencephalon  of  a  Full-term  Human 

Fetus.     (Cunningham.) 

gracile  and  cuneate  nuclei.  The  substance  of  Rolando  is  increased  and  be- 
comes rounded,  reaching  almost  to  the  surface  of  the  bulb  on  each  side,  only 
a  small  tract  of  longitudinal  fibers  of  the  root  of  the  fifth  nerve  intervening. 
There  is  a  great  increase  of  the  reticular  formation  around  the  central  canal. 
Then  at  the  ventral  or  anterior  aspect  the  decussation  of  the  pyramids  begins. 
By  this  crossing  over  of  the  fibers,  the  tip  of  the  gray  anterior  cornu  is  cut  off 
from  the  rest  of  the  gray  matter.  The  central  canal  is  pushed  farther  toward 
the  posterior  surface,  first  of  all  by  the  decussation  of  the  anterior  pyramids 
just  mentioned,  and  later  on,  i.e.,  above,  by  another  decussation  of  more 
dorsal  fibers.  These  fibers  of  the  second  decussation  as  they  cross  form  a 
median  raphe  and  also  help  to  break  up  the  remaining  gray  matter  into  what 
is  called  a  reticular  formation.  These  fibers  arise  from  the  nuclei  of  the  funic- 


ANATOMICAL    STRUCTURE 


609 


6lO  THE   NERVOUS    SYSTEM 

ulus  gracilis  and  funiculus  cuneatus  of  either  side,  and  they  are  looked  upon 
as  a  sensory  decussation. 

The  olivary  bodies  extend  forward  almost  to  the  level  of  the  pons.  They 
consist  of  both  cells  and  fibers.  The  cellular  matter  consists  of  a  plicated 
thinnish  layer  of  small  nerve  cells,  folded  upon  itself  in  the  form  of  a  loop, 
with  the  ends  turned  inward  and  slightly  dorsal,  figure  381,  O.  The  gray 
loop  is  filled  with  and  covered  by  white  fibers. 

Internal  to  the  olivary  body  on  either  side  are  two  small  masses  of  gray 
matter,  one  more  ventral  to  the  other,  called  accessory  olives,  external  and 


FIG.  380. — Anterior  or  Dorsal  Section  of  the  Medulla  Oblongata  in  the  Region  of 
the  Superior  Pyramidal  Decussation.  a.m.f.,  Anterior  median  fissure;  f,a.,  superficial 
arciform  fibers  emerging  from  the  fissure;  py,  pyramid;  n.ar.,  nuclei  of  arciform  fibers;  /.a., 
deep  arciform  becoming  superficial;  o,  lower  end  of  olivary  nucleus;  n.l.,  nucleus  lateralis; 
f.r.,  formatio  reticularis;/.a.2,  arciform  fibers  proceeding  from  the  formatio  reticularis; 
£.,  substantia  gelatinosa  of  Rolando;  a.V.,  ascending  root  of  fifth  nerve;  n.c.,  nucleus 
cuneatus;  n.c/,  external  cuneate  nucleus;  n.g.,  nucleus  gracilis;  /.#.,  funiculus  gracilis; 
p.m.f.,  posterior  median  fissure;  c.c.,  central  canal  surrounded  by  gray  matter,  in  which 
are  n.XI.,  nucleus  of  the  spinal  accessory,  and  n.XII.,  nucleus  of  the  hypoglossal;  s.d., 
superior  pyramidal  decussation.  (Modified  from  Schwalbe.) 

internal,  and  on  the  surface  of  the  anterior  pyramid  on  either  side  a  small 
mass  of  gray  matter,  external  arcuate  nucleus;  laterally  another  mass  of  the 
same  material,  the  representative  of  the  lateral  nucleus  of  the  cord,  is  seen, 
viz.,  the  antero-lateral  nucleus,  which  gives  origin  to  the  spinal  accessory 
nerve. 

It  will  be  necessary  to  follow  as  shortly  as  possible  the  fibers  of  the  spinal 
cord  upward  into  the  bulb  and  beyond. 

Tracts  Through  the  Medulla.— The  crossed  and  direct  pyramidal  tracts 
have  already  been  described.  Nothing  definite  is  known  of  the  antero-lateral 
descending  tracts.  The  direct  cerebellar  tracts  pass  laterally  into  the  resti- 


CONNECTIONS    OF    THE   BULB  6ll 

form  bodies  and  go  to  the  cerebellum.  The  antero-lateral- ascending 
tracts  (Gowers)  appear  to  have  the  same  destination,  but  pass  indirectly  into 
the  cerebellum  by  way  of  the  superior  medullary  velum;  some  of  the  fibers 
probably  pass  upward  to  higher  centers.  The  fibers  of  the  tracts  of  Goll 
and  Burdach,  of  the  cord,  end  in  the  nuclei  of  the  funiculus  gracilis  and  funic- 
ulus  cuneatus,  respectively;  at  any  rate,  ascending  degeneration  of  these 
columns  cannot  be  traced  above  these  nuclei.  The  rest  of  the  fibers  of  the 
cord  appear  to  end  in  the  reticular  formation  of  the  bulb. 

TT.TZT.        .     ,    «       ,  n>c 


7*.  err. 


FIG.  381. — Section  of  the  Medulla  Oblongata  at  about  the  Middle  of  the  Olivary  Body. 
/./.a.,  Anterior  median  fissure;  n.ar.,  nucleus  arciformis;  p.,  pyramid;  XII.,  bundle  of 
hypoglassal  nerve  emerging  from  the  surface;  at  b,  it  is  seen  coursing  between  the  pyramid 
and  the  olivary  nucleus,  o.;  f.a.e.,  external  arciform  fibers;  n.L,  nucleus  lateralis;  a., 
arciform  fibers  passing  toward  restiform  body,  partly  through  the  substantia  gelatinosa, 
#.,  partly  superficial  to  the  ascending  root  of  the  fifth  nerve,  a,V.;  X.,  bundle  of  vagus  root 
emerging;  f.r.,  formatio  reticularis;  c.r.,  corpus  restiforme,  beginning  to  be  formed,  chiefly 
by  arciform  fibers,  superficial  and  deep;  n.c.,  nucleus  cuneatus;  n.g.,  nucleus  gracilis;  *, 
attachment  of  the  ligula;/.s.,  funiculus  solitarius;  n.X.,  Xn..',  two  parts  of  the  vagus  nucleus; 
n.  XII.,  hypoglossal  nucleus;  n.t.,  nucleus  of  the  funiculus  teres;  n.am.,  nucleus  ambiguus; 
r.,  raphe;  A.,  continuation  of  the  anterior  column  of  cord;  o',o",  accessory  olivary  nucleus; 
P.O.,  pedunculus  olivas.  (Modified  from  Schwalbe.) 

Connections  of  the  Bulb  with  the  Cerebrum  and  Cerebellum. — The 

pyramidal  tracts  connect  the  bulb  with  the  cerebrum;  and  the  direct  cere- 
bellar  and  the  antero-lateral  ascending  tract,  tract  of  Gowers,  connect  it  with 
the  cerebellum.  Other  connections  of  the  bulb  with  the  cerebrum  and  with 
the  cerebellum  are: 

i.  Fibers  from  the  nucleus  gracilis  and  nucleus  cuneatus,  which,  as  we 
have  said,  are  the  endings  of  the  fibers  of  the  tracts  of  Goll  and  Burdach 
of  the  cord,  pass  in  sets  in  the  following  manner: 

a.  Internal  arcuate  fibers  pass  down  and  inward  to  the  opposite  side  in 
the  reticular  formation,  composing  in  part  the  superior  or  sensory  decussation, 


6l2  THE    NERVOUS    SYSTEM 

and  in  the  inter-olivary  region  enter  the  mesial  fillet,  which  passes  upward 
through  the  pons  to  end  about  the  cells  in  the  mid-brain  and  in  the  optic 
thalami.  These  fibers  are  probably  augmented  by  the  addition  of  fibers 
from  the  anterior  columns  of  the  cord,  and  by  fibers  arising  from  cells  in  the 
sensory  nuclei  of  the  cranial  nerves  ending  in  the  bulb. 

b.  External  arcuate  fibers,  after  decussating  in  the  same  way,  pass  out- 
ward superficially  over  the  anterior  pyramid  and  olivary  body,  reaching  the 
restiform  body  and  passing  to  the  side  of  the  cerebellum  opposite  to  their 
nuclei  of  origin.     These  fibers  appear  to  be  interrupted,  at  least  in  part,  in 
the  external  arcuate  nuclei.     They  connect  one  side  of  the  spinal  cord  with 
the  opposite  side  of  the  cerebellum  through  the  gracile  and  cuneate  nuclei. 

c.  Direct  lateral  fibers  pass  to  the  restiform  body  and  so  to  the  same  side 
of  the  cerebellum. 

2.  Fibers  from  the  olivary  body  pass  to  the  opposite  side  of  the  cerebellum 
through  the  reticular  formation  and  restiform  body. 

3.  Fibers  from  the  vestibular  nucleus  of  the  eighth  or  auditory  nerve  in 
the  floor  of  the  fourth  ventricle  pass  to  the  same  side  of  the  cerebellum. 

FUNCTIONS  OF  THE  MEDULLA. 

The  medulla  is  of  great  importance  in  the  physiological  economy  of 
the  body.  Its  functions  can  be  classified  in  three  groups.  First  should  be 
given  the  function  of  conduction  between  the  cord  and  the  higher  centers 
of  the  brain.  Second,  the  medulla  is  a  center  of  numerous  reflex  activities, 
especially  those  regulating  the  vital  functions  of  the  body,  such  as  respiration, 
circulation,  and  the  like.  A  third  function  of  centers  in  the  medulla  is  that 
of  automatic  activity. 

The  Medulla  as  a  Conducting  Path.— The  medulla  is  the  pathway 
of  all  ascending  and  descending  nerve  impulses  between  the  spinal  cord  and 
most  of  the  peripheral  sensory  and  motor  apparatus  on  the  one  hand,  and  the 
cerebellum  and  the  cerebral  centers  on  the  other.  These  conducting  paths 
are  described  in  the  tracts  that  have  already  been  discussed  at  some  length. 
They  are  represented  graphically  in  the  diagrams,  figures  379,  382,  and  388. 

Reflex  Centers  of  the  Medulla. — The  larger  number  of  the  cranial 
nerves,  as  we  shall  presently  see,  take  their  origin  from  the  medulla  and 
pons.  Some  of  these  nerves  have  both  sensory  and  motor  roots,  while 
others  are  either  exclusively  motor  or  sensory.  A  large  percentage  of  the 
afferent  or  sensory  impulses  that  enter  the  medulla  produce  reflex  effects 
on  the  motor  nuclei  so  richly  represented  in  the  medulla.  The  nuclei,  or 
centers,  regulating  some  of  the  most  important  functions  of  the  body  are 
among  those  in  this  group.  When  certain  of  these  centers  are  interfered 
with,  death  follows. 

Life  may  continue  when  the  spinal  cord  is  cut  away  in  successive  portions 


REFLEX    CENTERS    OF    THE    MEDULLA 


613 


from  below  upward  as  high  as  the  point  of  origin  of  the  phrenic  nerves  from 
the  cervical  cord.  In  amphibia,  the  brain  has  been  all  removed  from  above, 
and  the  cord  removed  as  far  as  the  medulla  oblongata  from  below;  yet  so  long 
as  the  medulla  oblongata  was  left  intact,  respiration  and  life  were  maintained. 
But  if  the  medulla  oblongata  is  wounded,  particularly  if  it  is  wounded  in  its 
central  part  opposite  the  origin  of  the  vagi,  the  respiratory  movements  cease, 


FIG.  382. — Diagram  of  Ascending  Conduction  Paths  from  the  Cord  through  the  Medulla 
and  the  Thalamus  to  the  Cerebral  Cortex.     (Cunningham.) 


and  the  animal  dies  from  asphyxiation.     This  effect  ensues  even  when  all 
parts  of  the  nervous  system  except  the  medulla  oblongata  are  left  intact. 

Injury  and  disease  in  men  are  accompanied  by  the  same  nerve  disturb- 
ances as  are  exhibited  by  these  experiments  on  animals.  Numerous  in- 
stances are  recorded  in  which  injury  to  the  medulla  oblongata  has  produced 
instantaneous  death;  and,  indeed,  it  is  through  injury  to  it,  or  of  the  part  of 
the  cord  connecting  it  with  the  origin  of  the  phrenic  nerves,  that  death  is 
commonly  produced  in  fractures  attended  by  sudden  displacement  of  the 
upper  cervical  vertebrae. 


614  THE    NERVOUS    SYSTEM 

The  majority  of  the  medullary  centers  are  reflex  centers  simply,  and  are 
stimulated  by  afferent  or  by  voluntary  impulses.  Some  of  them  are  auto- 
matic centers  and  are  capable  of  sending  out  efferent  impulses  without  pre- 
vious stimulation  by  afferent  or  by  voluntary  impulses.  The  automatic 
centers  are,  however,  normally  influenced  by  reflex  or  by  voluntary 
impulses. 

Some  of  these  reflex  centers  which  are  bilateral  are:  i.  Centers  for  the 
movements  of  deglutition.  The  medulla  oblongata  contains  in  the  motor 
nuclei  of  the  ninth  and  tenth  nerves  the  centers  whence  are  derived  the  motor 
impulses  enabling  the  muscles  of  the  palate,  pharynx,  and  esophagus  to  pro- 
duce the  successive  co-ordinated  and  adapted  movements  necessary  to  the 
act  of  deglutition,  page  339.  This  is  proved  by  the  persistence  of  the  act  of 
swallowing  in  some  of  the  lower  animals  after  destruction  of  the  cerebral 
hemispheres  and  cerebellum;  its  existence  in  anencephalous  monsters, 
figure  400;  and  by  the  complete  arrest  of  the  power  of  swallowing  when 
the  medulla  oblongata  is  injured  in  experiments. 

2.  Centers  for  the  combined  muscular  movements  of  sucking,  the  nerves 
concerned  being  the  facial  for  the  lips  and  mouth,  the  hypoglossal  for  the 
tongue,  and  the  inferior  maxillary  division  of  the  fifth  for  the  muscles  of  the 
jaw. 

3.  Centers  for  the  secretion  of  saliva,  which  have  been  already  mentioned, 
page  349- 

4.  Centers  for  vomiting,  page  376. 

5.  Centers  for  coughing,  which  is  a  reflex  act  quite  independent  of  the 
respiratory  act. 

6.  Centers  for  the  dilatation  of  the  pupil,  the  fibers  from  which  pass  out 
through  the  spinal  cord  in  the  two  upper  dorsal  nerves  into  the  cervical 
sympathetic. 

7.  The  respiratory  center  of  the  medulla  has  already  been  discussed  as 
regards  its  automatic  action.     It  is  only  necessary  to  repeat  here  that 
although  it  is  automatic  in  its  action,  being  capable  of  direct  discharge  of 
respiratory  impulses  with  no  other  stimulus  than  the  condition  of  the  blood 
circulating  within  it,  yet  it  is  constantly  reflexly  influenced  by  afferent  im- 
pulses.    The  respiratory  center  has  been  proven  to  be  bilateral.     It  also 
consists  of  an  inspiratory  part  and  of  an  expiratory  part.     The  center  is 
influenced  by  voluntary  impulses,  but  one  cannot  voluntarily  control  this 
center  to  the  point  of  death.     The  vagus  influence  is  probably  the  most 
constant  of  those  stimulating  the  respiratory  center.     But  the  respiratory 
reflexes  are  taking  place  constantly  in  response  to  afferent  impulses  flowing 
into  the  medulla  from  numerous  other  sensory  nerves  over  the  entire  body. 

8.  The  Cardio-inhibitory  Centers. — The  medulla  contains  the  centers 
which  maintain  the  proper  rhythm  of  the  heart,  these  centers  acting  through 
the  vagus  fibers.     These  terminate  in  a  local  intrinsic  mechanism  which  has 


THE    PONS   VAROLII  615 

been  already  discussed.  It  is  claimed  that  the  center  can  be  stimulated 
directly,  as  by  the  condition  of  the  blood  circulating  within  it.  It  is  con- 
stantly exerting  a  tonic  influence  over  the  heart,  which  is  the  chief  reason  for 
considering  it  an  automatic  center.  But  the  cardio-inhibitory  center  is 
primarily  a  reflex  center.  Sensory  or  afferent  impulses  arriving  over  the 
sensory  paths  in  the  vagus  itself,  by  abdominal  paths  through  the  sympa- 
thetic, and  through  cutaneous  nerves,  are  constantly  causing  reflex  discharges 
of  inhibitory  impulses  from  this  center. 

9.  Accelerator  centers  for  the  heart  are  present  in  .the  medulla.     They 
are  reflexly  stimulated  by  sensory  impulses  arising  from  the  same  general 
source  as  in  the  preceding  center. 

10.  Vaso-motor  centers  which  control  the  unstriped  muscle  of  the  arteries, 
are  also  situated  in  the  medulla.     The  nerve  cells  constituting  the  center 
are  under  the  constant  influence  of  nerve  impulses  flowing  in  from  the  sensory 
and  motor  structures  throughout  the  whole  body.     The  reflexes  produced  by 
the  afferent  impulses  bring  about  the  variations  in  vaso-motor  tone  that  not 
only  regulate  the  general  vascular  responses  of  the  body,  but  control  and  co- 
ordinate the  local  changes  in  the  size  of  the  blood  vessels. 

11.  Centers  for  the  secretion  of  sweat  exist  in  the  medulla.     The  medul- 
lary centers  control  the  subsidiary  spinal  sweat  centers.     They  may  be 
excited  unequally  so  as  to  produce  unilateral  sweating. 

The  reflex  medullary  centers  described  above  are  comparable  to  the  spinal 
reflex  centers  previously  described.  If  the  medulla  were  completely  isolated 
from  the  higher  cerebral  centers,  and  the  spinal  cord  removed  with  the  ex- 
ception of  those  paths  which  are  necessary  to  maintain  respiration,  these 
medullary  reflex  centers  would  be  able  to  co-ordinate  afferent  impulses  in  the 
same  general  way  that  isolated  segments  of  the  cord  do.  In  the  living  body, 
however,  the  medullary  centers  are  under  the  influence  of  changes  going  on 
in  regions  of  the  nervous  system  both  above  and  below,  changes  which  con- 
stantly influence  the  details  of  the  reactions.  The  activities  are  unconscious 
reflexes  in  the  same  sense  that  the  motor  reflexes  of  the  spinal  cord  are  un- 
conscious and  machine-like.  The  main  difference  is  one  of  complexity  and 
not  of  kind. 

The  Pons  Varolii. — The  pons  Varolii  is  generally  spoken  of  as  a  great 
commissure  of  fibers;  of  fibers  which  connect  the  two  halves  of  the  cerebellum 
and  which  connect  the  bulb  and  spinal  cord  with  the  cerebellum.  It  must  not 
be  forgotten  that  the  pons  contains  several  smaller  collections  of  nerve  cells. 
Sections  reveal  the  following  parts  or  structures,  beginning  with  the  anterior 
or  ventral  surface. 

i.  Transverse  or  commissural  fibers  connect  one  side  of  the  cerebellum 
with  the  other  through  the  middle  peduncle.  These  fibers  connect  the  cere- 
bellar  cortex  with  the  cells  of  the  pontine  nuclei;  some  are  afferent,  some 
efferent;  some  end  in  the  gray  matter  of  the  pons  on  the  same  side  near  the 


6i6 


THE    NERVOUS    SYSTEM 


ventral  surface;  others  cross  to  the  opposite  side  of  the  pons  and  then  become 
longitudinal,  passing  on  to  the  tegmentum. 

2.  Fibers  longitudinal  in  direction  are  arranged  in  larger  or  smaller  bun- 
dles and  are  separated  by  gray  matter.     Most  of  these  fibers  are  pyramidal 
fibers  which  pass  down  to  the  pyramids  of  the  medulla. 

3.  The  dorsal  portion  of  the  pons  is  made  up  to  a  considerable  extent  of 
the  reticular  formation  of  the  tegmental  region  together  with  one  or  two 
distinct  bundles  of  longitudinal   fibers.     The   chief  longitudinal  bundle, 
situated  at  the  junction  of  the  ventral  two- thirds  with  the  dorsal  third,  is  the 


FIG.  383. — Scheme  to  show  the  connections  of  the  Posterior  Longitudinal  Bundle. 
(Cunningham,  modified  from  Held.) 


fillet,  including  a,  the  larger  mesial  fillet,  a  sensory  tract  previously  described 
arising  in  the  gracile  and  cuneate  nuclei,  and  b,  the  lateral  fillet,  an  auditory 
tract,  see  figure  414.  The  second,  the  posterior  longitudinal  bundle,  is  situated 
on  each  side  of  the  mid-line,  just  internal  to  the  mesial  fillet.  Some  of  the 
connections  of  the  posterior  longitudinal  bundle  are  shown  in  figure  383. 

4.  In  the  upper  part  of  the  pons  there  is  a  mass  of  gray  matter  containing 
pigment,  the  locus  ceruleus;  and  in  the  back  part  a  second  mass  of  gray  matter, 
the  superior  olive,  which  is  connected  with  the  auditory  conduction  path, 
figure  414. 


THE    CEREBELLUM  617 

THE  CEREBELLUM. 

The  cerebellum  is  a  large  division  of  the  brain,  located  just  beneath  the 
cerebrum  and  behind  the  medulla  and  pons.  It  is  connected  with  the  rest 
of  the  brain  by  three  peduncles  on  each  side:  the  superior,  the  middle,  and  the 
inferior  peduncle,  figure  384. 

The  cerebellum  is  composed  of  white  and  gray  matter,  the  latter  being 
external  as  in  the  cerebrum,  and,  like  it,  infolded  so  that  a  larger  area  may  be 
contained  in  a  given  space.  The  convolutions  of  the  gray  matter,  however, 


FIG.  384. — Cerebellum  in  Section  and  Fourth  Ventricle,  with  the  Neighboring  Parts. 
I,  Median  groove  of  fourth  ventricle,  ending  below  in  the  calamus  scriptorius,  with  the 
longitudinal  eminences  formed  by  the  fasciculi  teretes,  one  on  each  side;  2,  the  same  groove, 
at  the  place  where  the  white  streaks  of  the  auditory  nerve  emerge  from  it  to  cross  the  floor 
of  the  ventricle;  3,  inferior  crus  or  peduncle  of  the  cerebellum,  formed  by  the  restiform 
body;  4,  posterior  pyramid;  above  this  is  the  calamus  scriptorius;  5,  superior  crus  of 
cerebellum,  or  processus  e  cerebello  ad  cerebrum  (or  ad  testes);  6,  6,  fillet  to  the  side  of  the 
crura  cerebri;  7,  7,  lateral  grooves  of  the  crura  cerebri;  8,  corpora  quadrigemina.  (From 
Sappey,  after  Hirschfeld  and  Leveille.) 


are  arranged  after  a  different  pattern,  as  shown  in  figure  385.  Besides  the 
gray  substance  on  the  surface,  there  is,  near  the  center  of  the  white  substance 
of  each  hemisphere,  a  small  capsule  of  gray  matter  called  the  corpus  dentatum, 
figure  385,  resembling  very  closely  the  corpus  dentatum  of  the  olivary  body  of 
the  medulla  oblongata. 

If  a  section  be  taken  through  the  gray  matter  of  the  cerebellum,  it  will  be 
found  to  be  composed  of  two  layers,  an  outer,  or  molecular,  and  an  inner,  or 
granular,  layer.  Each  of  these  layers  contains  a  large  number  of  peculiar- 
shaped  nerve  cells  and  very  rich  plexuses  of  nerve  fibers.  Recent  studies 
of  the  cortex  of  the  cerebellum  by  modern  methods  have  revealed  a  most 
complex  and  beautiful  arrangement  of  the  parts. 


6i8 


THE    NERVOUS    SYSTEM 


The  General  Structure  of  the  Cerebellum.— The  molecular  layer 
of  the  cerebellum  contains  several  peculiar  types  of  nerve  cells,  of  which  may 
be  specially  mentioned  Purkinje's  cells  and  the  basket  cells.  The  cells  of 
Purkinje  lie  along  the  internal  margin  of  the  molecular  layer,  being,  in  fact, 
practically  at  the  boundary  of  the  molecular  and  granular  layers.  They 
measure  40  to  60  /*  in  diameter,  and  have  large,  round  nuclei.  Each  cell 
gives  off  an  enormous  number  pf  branching  dendrites,  which  run  up  toward 
the  surface  of  the  cerebellum  in  the  shape  of  a  bush. 

The  cells  of  Purkinje  give  off  at  their  deeper  surface  an  axone  which 
runs  down  into  the  white  matter  of  the  cerebellum.  Recurrent  collaterals 
occur. 


FIG.  385. — Outline  Sketch  of  a  Section  of  the  Cerebellum,  Showing  the  Corpus 
Dentatum.  The  section  has  been  carried  through  the  left  lateral  part  of  the  pons,  so  as  to 
divide  the  superior  peduncle  and  pass  nearly  through  the  middle  of  the  left  cerebellar 
hemisphere.  The  olivary  body  has  also  been  divided  longitudinally  so  as  to  expose  in 
section  its  corpus  dentatum.  cr,  Crus  cerebri;/,  fillet;  q,  corpora  quadrigemina;  sp,  superior 
peduncle  of  the  cerebellum,  divided;  mp,  middle  peduncle  or  lateral  part  of  the  pons  Varolii, 
with  fibers  passing  from  it  into  the  white  stem;  av,  continuation  of  the  white  stem  radiating 
toward  the  arbor  vitse  of  the  folia;  o,  olivary  body  with  its  corpus  dentatum;  p,  anterior 
pyramid.  (Allen  Thomson.) 

Lying  in  the  molecular  layer,  somewhat  external  to  the  Purkinje  cells, 
are  the  cells  of  the  type  known  as  basket  cells.  These  cells  have  a  number  of 
dendrites;  they  also  send  out  an  axone  which  runs  parallel  to  the  surface  of 
the  cortex  and  gives  off  numerous  collaterals  in  its  course  that  form  baskets 
around  the  cell  bodies  of  the  Purkinje  cells,  figure  386,  ZK. 

The  granular  layer  contains  a  large  number  of  very  small  granule-like 
cells  that  Golgi  was  the  first  to  show  are  really  nerve  cells.  They  are  only 
about  5  //  in  diameter,  and  they  have  a  number  of  short  dendrites  which  end  in 
clubbed  extremities.  They  give  off  a  very  slender  axis-cylinder  process  or 
axone  which  runs  up  into  the  superficial  part  of  the  molecular  layer  and 
there  divides  in  a  T-shaped  fashion,  the  fibers  run  parallel  to  the  surface 
of  the  convolution  and  pass  in  between  the  branches  of  the  cells  of  Purkinje. 

The  white  substance  of  the  cerebellum  consists  of  nerve  fibers,  which  are 


PATHS  THROUGH  THE  CEREBELLAR  CORTEX 

of  three  kinds:  i.  Descending  fibers,  that  are  made  up  of  the  axis-cylinders 
of  the  cells  of  Purkinje,  carrying  impulses  down  from  the  cerebellar  cortex. 
2.  Ascending  fibers,  which  pass  into  the  granular  layer,  and  there  end  in  a 
number  of  very  short,  finely  divided  brushes  of  fibers  presenting  a  mossy 
appearance,  so  that  these  are  known  as  the  mossy  fibers.  These  connect 
with  the  granular  cells  of  this  layer.  3.  Ascending  fibers,  which  pass  up 


FIG.  386.— Transverse  Section  Through  a  Cerebellar  Folium  (after  Kolliker).  Treated 
by  the  Golgi  method.  P,  Axone  of  cell  of  Purkinje;  F,  moss  fibers;  K  and  K'y  fibers  from 
white  core  of  folium  ending  in  molecular  layer  in  connection  with  the  dendrites  of  the  cells 
of  Purkinje;  M,  simple  cell  of  the  molecular  layer;  GR,  granule  cell;  GR1,  axones  of  granule 
cells  in  molecular  layer  cut  transversely;  M',  basket  cells;  ZK,  basket  work  around  the  cells 
of  Purkinje;  GL,  neuroglia  cell;  N,  axone  of  an  association  cell. 

through  the  granular  into  the  molecular  layer  and  there  break  up  into  a  fine 
network  which  interlaces  with  the  dendritic  branches  of  the  cells  of  Purkinje. 

Paths  through  the  Cerebellar  Cortex.— It  will  be  seen  that  the  ar- 
rangements for  the  transmission  and  diffusion  of  nerve  impulses  and  for  the  co- 
operation of  different  cells  are  extremely  complicated  and  delicate.  It  is  not 
possible  to  indicate  absolutely  by  any  schema  the  course  of  fibers  and  the 
course  of  impulses  through  the  cerebellum,  but  approximately  it  is  some- 
what like  that  in  the  accompanying  figure  387. 

Impulses  pass  up  along  the  ascending  fibers  to  the  granular  cells  by  way  of 
the  direct  cerebellar,  the  fibers  of  the  gracile  and  of  the  cuneatus,  from  the 


62O 


THE    NERVOUS    SYSTEM 


restiform  body,  etc.  These  cells,  being  stimulated,  send  the  impulses  by  their 
axis-cylinders  to  the  molecular  layer,  and  through  their  T-shaped  divisions  to 
the  dendrites  of  the  cells  of  Purkinje.  Thence  an  impulse  is  sent  out  by  the 
axis-cylinder  process  of  this  cell.  Other  ascending  impulses  are  brought  up 
by  those  fibers  which  pass  directly  to  the  molecular  layer  and  send  their  ter- 
minals winding  around  the  dendrites  of  the  cells  of  Purkinje.  Prob- 
ably impulses  pass  up  also  through  the  ascending  fibers  which  affect  the 
basket  cells,  and,  through  them  and  their  basket-like  terminals,  the  cells  of 
Purkinje.  Purkinje  cells  send  cerebeilar  motor  fibers  to  the  nucleus  dentatus 


Stellate 


molecular 
layer 

'""-^^ 

granule 

granule 


FIG.  387. — A,  Afferent  fiber  to  basket  (stellate)  cell;  B,  neuraxone  of  Purkinje  cell;  C, 
afferent  fiber  to  Purkinje  cell;  D,  afferent  (mossy)  fiber  to  granule  cell. 

cerebelli  and  through  the  superior  peduncles  to  the  red  nucleus  and  the 
thalamus,  and  to  the  ventro-lateral  descending  tract  of  the  cord,  to  end 
about  the  anterior  horn  cells. 

Functions  of  the  Cerebellum. — With  the  exception  of  its  middle 
lobe,  the  cerebellum  is  itself  insensible  to  irritation  and  may  be  all  cut  away 
without  eliciting  signs  of  pain  (Longet).  Its  removal  or  disorganization  by 
disease  is  also  generally  unaccompanied  by  loss  or  disorder  of  sensibility; 
animals  from  which  it  is  removed  can  smell,  see,  hear,  and  feel  pain,  to  all 
appearances,  as  perfectly  as  before  (Flourens;  Magendie).  It  cannot,  there- 
fore, be  regarded  as  a  principal  organ  of  sensation.  Yet  if  any  of  its  crura 
be  touched,  pain  is  indicated;  and,  if  the  restiform  tracts  of  the  medulla 
oblongata  be  stimulated,  the  most  acute  suffering  appears  to  be  produced. 

These  phenomena  may  properly  be  ascribed  to  the  activity  of  the  cerebral 
cortex,  since  the  number  of  collaterals  on  the  fibers  that  pass  to  cerebeilar 
tracts  is  very  great,  and  impulses  arising  from  their  stimulation  may  reach 
the  sensorium  by  paths  other  than  through  the  cerebellum. 


FUNCTIONS    OF    THE    CEREBELLUM 


621 


The  experiments  of  Longet  and  many  others  agree  in  supporting  the  view 
that  no  stimulation  of  the  cerebellar  cortex  leads  to  localized  muscular  con- 
tractions. In  other  words,  there  is  no  localization  in  the  cerebellar  cortex  as 
in  the  cerebrum,  the  cerebellum  apparently  acting  as  a  whole.  If  the  cere- 
bellum be  removed,  as  was  done  by  Flourens  and  numerous  later  physiologists, 
a  very  profound  disturbance  in  motor  functions  occurs.  With  the  removal 


J^N^f  Cranial  Nerve 
(Vestibular) 


¥IG.  388.— Scheme  of  Principal  Ascending  Cerebrospinal  (black)  and  Cerebellar   (red) 
Conduction  Paths.     (Modified  from  Hardesty  in  Morris'  Anatomy.) 

of  the  superficial  layers  of  the  cerebellum,  in  pigeons  particularly,  there  is 
increasing  feebleness  and  lack  of  harmony  of  the  muscles  concerned  in  lo- 
comotion. When  the  entire  organ  is  cut  away  in  pigeons  they  lose  the  power 
of  walking,  flying,  and  of  standing  in  the  usual  erect  way.  Their  power  of 
preserving  equilibrium  is  lost,  the  most  characteristic  feature.  Birds  do  not 
remain  in  a  state  of  stupor,  but  attempt  to  carry  out  the  usual  muscular  activi- 


622  THE    NERVOUS    SYSTEM 

ties.  If  a  pigeon  is  laid  on  its  back  it  cannot  recover  its  erect  position,  though 
it  make  motions  to  do  so.  If  set  on  its  feet  it  will  fall  to  one  side  or  the  other, 
and  is  not  able  to  hold  its  head  in  the  customary  position.  The  endeavors  of 
the  animal  to  maintain  its  balance  are  insecure  and  uncertain,  resembling 
the  lack  of  muscular  control  of  a  drunken  man. 

Such  an  animal  does  not  lose  the  power  of  perceiving  sensations,  nor  of 
making  voluntary  efforts,  as  it  will  endeavor  to  avoid  the  blow  that  is 
threatened. 

The  experiments  afford  the  same  results  when  repeated  on  all  classes  of 
animals;  and  from  them  and  the  others  before  referred  to,  Flourens  inferred 
that  the  cerebellum  belongs  neither  to  the  sensory  nor  the  intellectual  ap- 
paratus; and  that  it  is  not  the  source  of  voluntary  movements,  although  it  be- 
longs to  the  motor  apparatus,  but  is  the  organ  for  the  co-ordination  of  the 
voluntary  movements,  or  for  the  excitement  of  the  combined  action  of  muscles. 

Such  evidence  as  can  be  obtained  from  cases  of  diseases  of  this  organ 
confirms  the  view  taken  by  Flourens;  and,  on  the  whole,  it  gains  support  from 
comparative  anatomy — animals  whose  natural  movements  require  most 
frequent  and  exact  combinations  of  muscular  contractions  being  those  whose 
cerebella  are  most  developed  in  proportion  to  the  spinal  cord. 

We  must  remember,  too,  that  the  cerebellum  is  connected  with  the  pos- 
terior columns  of  the  cord  through  the  cuneate  and  gracile  nuclei  as  well  as 
with  the  direct  cerebellar  tract,  all  of  which  probably  convey  to  the  middle 
lobe  muscular  sensations.  It  is  also  connected  with  the  auditory  nerves  and 
bulb  by  the  internal  and  external  arcuate  fibers;  and  with  the  tegmentum 
through  the  red  nuclei.  Its  connection  with  the  efferent  tracts  from  the 
different  cerebral  lobes  through  the  pons  is  also  highly  important.  Move- 
ments of  the  eyes  also  occur  on  direct  stimulation  of  the  middle  lobe.  It 
seems,  therefore,  to  be  connected  in  some  way  with  all  of  the  chief  sensory 
impulses  which  have  to  do  with  the  maintenance  of  the  equilibrium,  and  is 
generally  included  in  the  nervous  apparatus  which  is  supposed  to  govern 
this  function  of  our  bodies. 

Foville  supposed  that  the  cerebellum  is  the  organ  of  muscular  sense,  i.e., 
the  organ  by  which  the  mind  acquires  that  knowledge  of  the  actual  state 
and  position  of  the  muscles  which  is  essential  to  the  exercise  of  the  will  upon 
them;  and  it  must  be  admitted  that  all  the  facts  just  referred  to  are  as  well 
explained  on  this  hypothesis  as  on  that  of  the  cerebellum  being  the  organ  for 
combining  movements.  A  harmonious  combination  of  muscular  actions 
must  depend  as  much  on  the  capability  of  appreciating  the  condition  of  the 
muscles  with  regard  to  their  tension,  and  to  the  force  with  which  they  are 
contracting,  as  on  the  power  which  any  special  nerve  center  may  possess  of 
exciting  them  to  contraction.  And  it  is  because  the  power  of  such  harmonious 
movement  would  be  equally  lost,  whether  the  injury  to  the  cerebellum  involved 
injury  to  the  seat  of  muscular  sense  or  to  the  center  for  combining  muscular 
actions,  that  experiments  on  the  subject  afford  no  proof  in  one  direction  more 
than  the  other. 


THE    MID-BRAIN  623 

Forced  Movements. — The  influence  of  each  half  of  the  cerebellum 
is  directed  to  muscles  on  the  opposite  side  of  the  body;  and  it  would  appear 
that,  for  the  right  ordering  of  movements,  the  action  of  its  two  halves  must  be 
always  mutually  balanced  and  adjusted.  For  if  one  of  its  crura,  or  if  the 
pons  on  either  side  of  the  middle  line,  be  divided,  so  as  to  cut  off  from  the 
medulla  oblongata  and  spinal  cord  the  influence  of  one  of  the  hemispheres 
of  the  cerebellum,  strangely  disordered  movements  ensue — forced  movements. 
The  animals  fall  down  on  the  side  opposite  to  that  on  which  the  crus  cerebelli 
has  been  divided,  and  then  roll  over  continuously  and  repeatedly;  the  rotation 
being  always  round  the  long  axis  of  their  bodies,  and  generally  from  the  side 
on  which  the  injury  has  been  inflicted.  The  rotations  sometimes  take  place 
with  much  rapidity;  as  often,  according  to  Magendie,  as  sixty  times  in  a  min- 
ute, and  may  last  for  several  days.  Similar  movements  have  been  observed 
in  men;  as  by  Serres  in  a  man  in  whom  there  was  apoplectic  effusion  in  the 
right  crus  cerebelli;  and  by  Belhomme  in  a  woman  in  whom  an  exostosis 
pressed  on  the  left  crus.  They  may,  perhaps,  be  explained  by  assuming  that 
the  division  or  injury  of  the  crus  cerebelli  produces  paralysis  or  imperfect  and 
disorderly  movements  of  the  muscles  of  the  opposite  side  of  the  body.  Such 
movements  cease  when  the  other  crus  cerebelli  is  divided;  but  probably  only 
because  the  paralysis  of  the  body  is  thus  made  almost  complete.  Other 
varieties  of  forced  movements  have  been  observed,  especially  those  named 
lt  circus  movement,"  when  the  animal  operated  upon  moves  round  and  round 
in  a  circle;  and  again  those  in  which  the  animal  turns  over  and  over  in  a  series 
of  somersaults.  Nearly  all  these  movements  may  also  result  on  section  of 
one  or  other  of  the  following  parts:  viz.,  medulla,  pons,  cerebellum,  corpora 
quadrigemina,  corpora  striata,  optic  thalami,  and  even,  it  is  said,  of  the 
cerebral  hemispheres.  But  these  structures  are  parts  that  involve  tracts 
used  in  the  co-ordination  of  complex  muscular  movements. 

THE  MID-BRAIN. 

The  mid-brain  includes  the  crura  cerebri  and  the  corpora  quadrigemina. 

The  Peduncles  of  the  Cerebrum,  or  Crus  Cerebri. — The  crura 
diverge  from  the  anterior  edge  of  the  pons  Varolii  and  pass  upward  on  either 
side  toward  the  cerebral  hemispheres.  At  their  anterior  termination  each 
of  them  appears  to  have  upon  its  dorsal  surface,  to  the  inner  and  outer  sides, 
respectively,  two  large  masses  of  gray  matter  which  have  been  already  spoken 
of,  viz.,  the  optic  thalamus  and  the  corpus  striatum.  The  crus  is  made  up 
of  two  principal  parts.  The  crusta  or  pes  is  in  the  ventral  position,  and  the 
tegmentum  in  the  dorsal  position.  The  two  are  separated  by  the  substantia 
nigra. 

The  pes  consists  of  longitudinal  fibers  which  pass  anteriorly  between  the 
optic  thalamus  and  the  posterior  part  (lenticular  nucleus)  of  the  corpus  stria- 


624 


THE    NERVOUS    SYSTEM 


turn.  In  this  situation  the  fibers  form  a  compact  mass.  This  constitutes 
the  internal  capsule,  and  that  portion  of  it  which  forms  the  angle  at  which 
the  fibers  are  bent  is  called  the  genu  of  the  capsule.  The  internal  capsule 
spreads  out  dorsally  in  the  corona  radiata.  The  fibers  thus  have  the  form 
of  a  fan  bent  upon  itself  as  they  rise  to  pass  into  the  cerebral  hemisphere. 
The  fibers  of  the  internal  capsule  are  connected  with  different  districts  of  the 
cerebral  cortex.  Briefly  the  connections  are,  a,  the  fronto-pontine  fibers  are 
in  the  anterior  limb  of  the  capsule;  b,  the  pyramidal  fibers  in  the  genu  and 
the  anterior  part  of  the  posterior  limb;  c,  the  temporo-pontine  fibers  in  the 


FIG.  389. — Diagram  of  the  Motor  Tract  as  Shown  in  a  Diagrammatic  Horizontal 
Section  through  the  Cerebral  Hemispheres,  Crura,  Pons,  and  Medulla.  Fr.,  Frontal  lobe; 
Oc.t  occipital  lobe;  AF.,  ascending  frontal,  AP.,  ascending  parietal  convolutions;  PCF., 
pre-central  fissure,  in  front  of  the  ascending  frontal  convolution;  FR.,  fissure  of  Rolando; 
IFF.,  inter-parietal  fissure,  a  section  of  crus  is  lettered  on  the  left  side;  SN.,  substantia 
nigra;  Py.,  pyramidal  motor  fiber  which  on  the  right  is  shown  as  continuous  lines  con- 
verging to  pass  through  the  posterior  limb  of  1C.,  internal  capsule  (the  knee  or  elbow  of 
which  is  shown  thus),  upward  into  the  hemisphere  and  downward  through  the  pons  to 
cross  the  medulla  in  the  anterior  pyramids.  (Gowers.) 

posterior  part  of  the  posterior  limb.  Fibers  connecting  the  optic  thalami 
and  corpora  striata  with  the  cerebral  cortex  also  run  in  the  capsule.  The 
pes,  internal  capsule,  and  the  corona  radiata  form  the  great  sensory  and  motor 
highway  to  and  from  the  cerebral  cortex. 

The  tegmentum  is  the  continuation  anteriorly  of  the  reticular  formation  of 
the  medulla.  It  ends  for  the  most  part  in  the  neighborhood  of  the  optic 
thalamus  and  in  the  parts  beneath.  The  tegmentum  of  either  side  is  sup- 
posed to  be  concerned  chiefly  with  afferent  impulses.  It  is  made  up  to  a  very 
considerable  extent  of  collections  of  gray  matter,  the  most  important  of  which 
are  the  substantia  nigra,  separating  the  pes  and  tegmentum,  and  the  nucleus 
ruber,  which  is  a  rounded  mass  situated  near  the  aqueduct  of  Sylvius.  The 


THE    FORE -BRAIN  625 

latter  serves  as  a  way-station  in  the  cerebello-cerebral  conduction  paths  and 
also  has  important  connections  with  the  spinal  cord.  The  locus  niger  ex- 
tends back  as  far  as  the  posterior  corpus  quadrigeminum.  Posteriorly,  the 
tegmentum  is  chiefly  reticular  in  structure. 

Corpora  Quadrigemina. — There  are  two  co'rpora  quadrigemina  on  each 
side,  the  superior  and  inferior.  They  form  prominences  on  the  dorsal 
surface  of  the  mid-brain,  dorsal  to  the  aqueduct  of  Sylvius.  The  inferior 
corpora  quadrigemina  receive  through  the  lateral  fillet  fibers  from  the 
cochlear  division  of  the  eighth  nerve,  figure  414.  They  are  closely  associated 
with  the  median  corpora  geniculata,  and,  like  these,  give  origin  to  fibers 
which  continue  the  auditory  conduction  path  upward  to  the  auditory  center. 
The  superior  corpus  quadrigeminum  receives  fibers  from  the  optic  nerve, 
the  mesial  fillet,  and  also  from  the  occipital  cortex,  as  will  be  more  fully  de- 
scribed later.  It  is  closely  associated  with  the  external  corpus  geniculatum. 
The  corpora  geniculata  also  form  reflex  centers  for  the  eye  muscles  in  the 
reflexes  that  result  in  the  adjustments  of  the  eye  to  vision  at  different  dis- 
tances. The  nervous  connections  of  these  nuclei  are  discussed  in  presenting 
the  optic  tract  and  the  auditory  tracts. 

THE  FORE-BRAIN. 

The  fore-brain  or  prosencephalon  consists  of  two  divisons,  the  thalamen- 
cephalon  or  thalami,  and  the  telencephalon,  or  corpora  geniculata,  corpora 
striata,  and  the  cerebral  hemispheres.  These  subordinate  divisions  will  be 
discussed  in  order. 

The  Thalami. — The  optic  thalami  are  oval  in  shape,  and  rest  upon  the 
crura  cerebri.  They  form  part  of  the  floor  of  the  lateral  ventricles  and  their 
inner  sides  bound  the  third  ventricle.  They  are  connected  by  a  transverse 
tract,  the  middle  commissure. 

Each  thalamus  has  several  collections  of  gray  matter,  forming  somewhat 
indistinctly  defined  masses  separated  by  white  fibers.  These  masses  of  gray 
matter  are  known  as  the  nuclei  of  the  thalamus,  which  are  six  in  number. 
They  are  called  the  anterior  nucleus,  the  median  nucleus,  the  lateral  nucleus, 
the  ventral  nucleus,  the  pulvinar,  and  the  posterior  nucleus.  The  corpora 
geniculata  are  also  closely  associated  with  the  optic  thalamus.  The  anterior 
nucleus  is  composed  of  large  nerve  cells  which  receive  the  terminations  of 
axones  of  cells  of  the  corpora  mammillaria  at  the  base  of  the  brain  (bundle 
of  Vicq  d'Azyr).  There  they  meet  the  fibers  of  the  fornix,  which  establish 
a  relation  between  this  tubercle  of  the  thalamus  and  the  hippocampal  con- 
volutions. The  median  nucleus  is  connected  by  its  axones  with  the  cortex 
of  the  island  of  Reil  and  the  second  and  third  frontal  convolutions.  The 
lateral  nucleus  is  quite  large  and  lies  against  the  internal  capsule,  into  which 
it  sends  fibers.  It  is  connected  with  the  central  convolutions  of  the  cortex. 


626  THE    NERVOUS    SYSTEM 

The  ventral  nucleus  lies  beneath  the  preceding;  it  is  relatively  small.  It  is 
connected  with  the  cortex  of  the  frontal  lobe  and  with  the  operculum,  the 
central  convolutions,  and  the  supramarginal  gyrus.  The  fifth  nucleus, 
known  as  the  pulvinar,  forms  the  posterior  tip  of  the  thalamus,  and  is  con- 
nected with  the  optic  tract,  see  figure  406.  The  posterior  nucleus,  lying  just 
below  the  pulvinar,  is  a  small  mass  and  is  connected  with  the  cortex  of  the 
interior  parietal  convolution.  The  cells  of  the  thalamus  are  thus  seen  to  be 
connected  with  a  large  area  of  the  cerebral  cortex.  The  axones  spread  out 
in  a  great  fan  in  the  corona  radiata,  the  thalamus  sending  more  fibers  to  the 
cortex  than  are  received  from  it. 

The  collections  of  nerve  cells  in  the  thalamus  are  shown  by  anatomical 
investigations  and  by  methods  of  physiological  degeneration  to  be  on  the 
pathway  of  ascending  or  afferent  nerve  tracts.  Large  masses  of  sensory 
fibers  pass  through  the  thalami,  the  majority  of  which  form  synapses  about 
the  nerve  cells  in  the  thalamus.  Even  in  those  cases  where  there  is  no  dis- 
tinct ending  of  the  nerve  fiber,  collaterals  are  given  off  which  establish  physio- 
logical connection  with  the  nuclei. 

The  thalamus  is  thus  closely  connected  with  large  areas  of  the  cortex. 
It  must  at  least  form  an  important  relay  station  in  all  those  activities  which 
involve  the  conscious  perception  of  sensory  stimuli  wherever  they  may 
arise.  Flechsig  even  claims  that  in  the  thalamus  there  are  definite  points 
of  sensory  localization  corresponding  to  every  sensory  point  in  the  periphery 
of  the  body  (including  the  special  senses).  The  thalamus  also  receives 
fibers  from  various  parts  of  the  cerebral  cortex,  thus  establishing  a  double 
relation  with  this  region. 

Owing  to  the  difficulty  of  those  operations  which  establish  isolation  of  the 
thalamus,  it  is  not  clear  to  what  extent  reflex  actions  may  take  place  through 
its  nuclei.  It  is  probable,  however,  that  extensive  co-ordinations  of  afferent 
impulses  may  be  mediated  by  the  nuclei  of  the  thalami.  Such  activities  as 
walking,  riding,  writing,  speaking,  etc.,  are  possibly  reflexes  co-ordinated 
through  the  thalami,  perhaps  with  the  assistance  of  the  medulla  in  the  case 
of  walking. 

Corpora  Geniculata. — The  corpora  geniculata  form  prominences  on 
each  side  of  the  peduncles  of  the  cerebrum  just  ventral  to  the  anterior  corpora 
quadrigemina.  There  are  two  on  each  side,  the  external  or  outer,  and  median 
or  inner.  The  external  corpus  geniculatum  is  at  the  side  of  the  crus  and 
appears  to  be  a  swelling  on  the  lateral  division  of  the  optic  tract  and  actually 
receives  terminations  of  the  optic  fibers,  thus  constituting  a  way-station  in 
the  optic  conduction  paths.  Similarly  the  median  appears  to  be  the  ter- 
mination of  the  median  division  of  the  optic  tract,  from  which  it  receives 
some  fibers,  figure  406,  but  it  is  more  intimately  connected  with  the  auditory 
tracts,  forming  a  way-station  between  the  lateral  fillet  and  the  auditory  cor- 
tical center,  figure  414. 


THE    CEREBRUM  627 

Corpora  Striata. — The  corpora  striata  are  situated  in  front  and  to 
the  outside  of  the  thalami,  partly  within  and  partly  without  the  lateral 
ventricles. 

Each  corpus  striatum  consists  of  two  parts:  An  intraventricular  portion, 
the  caudate  nucleus,  which  is  conical  in  shape,  with  the  base  of  the  cone  for- 
ward (this  consists  chiefly  of  gray  matter),  and  an  extraventricular  portion, 
the  lenticular  nucleus,  separated  from  the  other  portion  by  the  internal  cap- 
sule. The  lenticular  nucleus  is  shown  in  a  horizontal  section  of  the  hemi- 
sphere to  consist  of  three  parts,  the  two  internal  called  globus  pallidus  major 
and  minor,  and  the  outer  called  the  putamen. 

The  cells  of  the  corpora  striata  are  somewhat  evenly  distributed,  and  not 
grouped  in  nuclei.  Their  axones  pass  for  the  most  part  into  the  internal 
capsule.  It  is  doubtful  if  these  ganglia  have  any  direct  anatomical  relations 
with  the  cortex  of  the  brain,  but  they  are  intimately  connected  by  fibers  to  and 
from  the  thalami,  and  are  connected  with  the  substantia  nigra  (Flechsig). 
These  nuclei  are  developed  from  the  walls  of  the  embryonic  brain  tube  and 
are  probably  therefore  homologous  with  the  areas  of  the  cortex.  Their  lesion 
is  said  to  be  accompanied  by  disturbance  in  muscular  co-ordination.  Lesion 
of  the  left  lenticular  nucleus  is  said  to  cause  some  disturbance  in  the  power  of 
speech,  though  this  has  not  been  observed  in  the  case  of  the  right  nucleus. 
Lesions  of  the  corpora  striata  produce  disturbances  in  heat  regulation, 
causing  a  rise  of  body  temperature,  the  rise  amounting  to  as  much  as  2°  or  3° 
C.  in  the  rabbit.  The  rise  of  temperature  in  man  after  lesion  of  the  corpus 
striatum  on  one  side  is  said  to  be  chiefly  on  the  opposite  side  of  the  body 
(Kaiser). 

THE  CEREBRUM. 

That  portion  of  the  brain  which  is  concerned  with  all  intellectual  functions 
is  the  cerebrum  or,  more  strictly  speaking,  the  cerebral  cortex.  The  cerebral 
cortex  is  the  seat  of  those  activities  which  we  describe  as  intelligence — 
including  states  of  consciousness,  acts  of  idea  formation  and  volition,  and 
the  phenomenon  of  memory. 

The  cerebrum  includes  the  cerebral  cortex,  the  mass  of  fibers  connecting 
it  with  lower  portions  of  the  brain,  the  basal  nuclei  represented  by  the 
corpora  striata,  the  thalami,  etc.  The  structure  and  function  of  these 
basal  nuclei  have  already  been  given  briefly,  so  we  may  turn  our  attention 
now  to  the  cerebral  cortex. 

Structure  of  the  Cerebral  Cortex. — The  cerebral  cortex  forms  a  large 
part  of  the  mass  of  the  cerebrum,  in  fact  of  the  whole  brain.  Its  superficial 
appearance  presents  a  series  of  ridges  and  folds,  the  gyri  and  sulci.  For  gen- 
eral convenience  anatomists  have  divided  the  cerebral  cortex  into  five  lobes: 
the  frontal,  that  portion  in  front  of  the  fissure  of  Rolando  extending  down 


628 


THE   NERVOUS    SYSTEM 


to  the  Sylvian  fissure;  the  parietal,  extending  from  the  Sylvian  fissure  to  the 
parieto-occipital  fissure,  and  bounded  below  by  the  Sylvian  fissure;  the 
temporal  lobe,  just  ventral  to  the  parietal;  the  central  lobe,  or  island  of 


cauaso-marcf. 


FIG.  390. — Left  Hemisphere,  from  Without.     (After  Eberstaller.) 


FIG.  391. — The  Cerebrum,  from  Above.     (After  Eberstaller.) 

Reil;  and  the  occipital  lobe,  which  includes  the  posterior  portion  of  the 
cortex  behind  the  parieto-occipital  fissure.  And,  finally,  the  olfactory  and 
limbic  lobes  together  make  up  the  olfactory  division  of  the  brain.  For  the 


STRUCTURE  OF  THE  CEREBRAL  CORTEX 


629 


detailed  arrangements  of  the  cortical  gyri  and  sulci  the  reader  is  referred  to 
figures  390,  392  and  to  text-books  of  anatomy. 

In  a  transverse  section  of  the  cerebral  cortex  there  is  shown  an  external 
gray  layer  chiefly  composed  of  nerve  cells  and  an  internal  white  portion  of 
nerve  fibers.  The  folding  of  the  cortex  into  convolutions  increases  the  total 
mass  of  gray  matter  enormously. 

The  gray  or  cellular  external  part  of  the  cerebral  cortex  has  an  average 
thickness  of  about  3  mm.;  being  thin  in  the  occipital  and  frontal  region, 
2mm.,  and  thick  in  the  precentral,  4  mm.,  and  postcentral  convolutions. 

Several  types  of  nerve  cells  have  been  described  as  present  in  the  cortex, 
the  exact  type  and  relative  proportion  varying  somewhat  in  different  regions. 


FIG.  392. — Right  Hemisphere,  from  Within.     (After  Eberstaller.) 

The  typical  characteristic  cell,  however,  is  the  pyramidal  cell.  The  py- 
ramidal cell,  as  its  name  implies,  has  a  pear-shaped  cell  body  with  numerous 
protoplasmic  processes.  The  apex  of  the  cell  is  directed  toward  the  surface 
of  the  cortex,  and  supports  numerous  branches  which  extend  out  into  the 
adjacent  territory,  bringing  it  into  contact  with  a  relatively  large  number  of 
nerve  cells.  These  processes  are  dendritic  in  character.  The  base  of  the 
pyramidal  cell  always  has  a  single  axis-cylinder  process  which  is  directed 
down  into  the  white  matter,  and  which  in  some  cases  ultimately  finds  its 
course  through  the  corona  radiata  into  the  pyramids  below.  The  axis- 
cylinder  processes  give  off  collaterals  both  in  the  immediate  neighborhood 
of  the  cell  and  somewhat  deeper  along  its  course. 

In  the  superficial  layer  of  the  cortex  there  is  a  peculiar  type  of  small  cell, 
first  described  by  Cajal.  Most  of  these  cells  are  fusiform  in  shape,  with  the 
long  axis  parallel  to  the  surface  of  the  convolution.  They  give  off  usually  two 
axones  which  run  along  parallel  to  the  surface  and  send  down  numerous 
fine  collaterals  at  right  angles.  Another  form  of  Cajal  cell,  triangular  or 


630 


THE    NERVOUS   SYSTEM 


quadrangular  in  shape,  is  also  seen.  Both  forms  have,  as  a  rule,  more  than 
one  axone.  Their  collaterals  pass  in  a  horizontal  direction,  forming  a 
fine  band  of  fibers,  known  as  tangential  fibers.  A  third  type  of  cell  is  the 
fusiform  or  polymorphous.  Some  of  these  are  strictly  fusiform  in  shape  and 
lie  with  their  axes  parallel  to  the  surface  of  the  convolution.  They  give  off 
protoplasmic  processes  which  pass  down  toward  the  white  matter,  some  of 


PIG.  393- 


FIG.  394. 


393- — Typical  Pyramidal  Cell  from  the  Human  Cortex,  a,  Cell  body;  6,  main 
dendrites  with  gemmules;  c,  lateral  dendrites;  d,  axone  and  collaterals.  Only  a  small 
part  of  the  axone  is  shown.  (Bailey.) 

FIG.  394. — Showing  the  Stages  in  the  Development  of  a  Pyramidal  Cell.     (Ram6n  y 

Cajal.) 

them  turning  to  run  in  a  horizontal  direction.  The  fusiform  and  poly- 
morphous cells  are  grouped  in  the  same  layer. 

Besides  these  cells  \\e  find  scattered  through  the  cortex  a  considerable 
number  of  the  neuroglia  cells.  The  character  and  position  of  these  are 
shown  in  figure  395. 

The  general  arrangement  of  the  layers  of  the  cortex  is  described  very  dif- 
ferently by  the  various  authors.  It  is  not  uniform  in  the  different  parts  of  the 
brain.  The  simplest  and  most  representative  type,  however,  of  the  arrange- 


STRUCTURE  OF  THE  CEREBRAL  CORTEX 


63I 


ment  is  that  in  which  the  cortex  is  divided  into  four  layers.  The  outermost, 
or  superficial,  known  as  the  molecular  layer,  contains  relatively  few  cells. 
It  is  composed  of  neuroglia  tissue,  embedded  in  which  are  a  number  of  cells 
of  the  Cajal  types,  which  have  just  been  described.  There  are  also  in  this 
layer  many  neuroglia  cells.  In  the  superficial  part  of  the  layer  of  some  areas 


FIG.  395. — The  Principal  Constituent  Elements  of  the  Gray  Cortical  Layer  of  the  Anterior 
Cerebrum.     (After  Ramdn  y  Cajal.) 

of  the  cortex  are  many  tangential  fibers.  The  second  layer  is  composed  of 
small  pyramidal  cells.  In  parts  of  the  brain  there  are  here  interposed 
what  are  known  as  the  vertical  fusiform  cells.  The  third  layer  is  composed 
of  large  pyramidal  cells,  in  which,  however,  one  also  sees  many  small  py- 
ramidal cells.  The  fourth  layer  is  composed  of  the  fusiform  and  polymorphous 
cells,  beneath  which  is  the  white  substance.  This  arrangement  is  shown 


632  THE    NERVOUS    SYSTEM 

in  the  accompanying  figures,  395  and  397.  The  gray  matter  of  the  brain 
contains,  however,  not  only  these  layers  and  cells,  but  an  infinitely  rich 
mass  of  fibers,  which  can  be  shown  to  have  a  certain  definite  arrangement. 
Some  of  the  fibers  are  vertical,  passing  directly  up  to  the  most  superficial 
layers  of  cells;  others  have  a  horizontal  direction,  dividing  the  gray  matter 
into  different  layers.  These  layers  of  fibers  have  received  different  names. 
A  typical  arrangement  is  shown  in  figure  398.  The  most  conspicuous 
fibers  are  those  of  certain  large  triangular  or  pyramidal  cells. 

The  efferent  or  axone  fibers  from  the  cerebral  cortex  may  be  divided  into 
three  classes:     i,  the  projection  fibers,  which  descend  through  the  corona 


FIG.  396. — Scheme  of  Descending  Conduction  Pathways  from  the  Cerebrum  to  Lower 

Nerve  Centers. 

radiata  and  internal  capsule,  to  end  in  lower  centers;  2,  the  commissural 
fibers,  which  cross  to  the  opposite  cerebral  hemisphere,  chiefly  through  the 
corpus  callosum;  3,  the  association  fibers,  which  pass  in  bundles  beneath 
the  cortex,  to  end  in  other  regions  of  the  same  hemisphere. 

It  is  by  means  of  projection  fibers  and  collaterals  that  associations  are 
made  with  nerve  cells  in  the  thalamus,  tegmentum,  and  pons,  and  through 
the  latter  region  with  tracts  going  to  the  cerebellum. 

Weight  of  the  Brain  and  Cord. — The  brain  of  an  adult  man  weighs  from  48 
to  50  oz.,  about  1,550  grams,  or  about  2  per  cent,  of  the  body  weight.  It 
exceeds  in  absolute  weight  that  of  all  the  lower  animals  except  the  elephant 
and  whale.  Its  weight,  relatively  to  that  of  the  body,  is  exceeded  only  by  that 
of  a  few  small  birds,  and  some  of  the  smaller  monkeys. 

In  the  new-born  child  the  brain  (weighing  10  to  14  oz.)  is  about  10  per 
cent,  of -the  total  body  weight.  At  the  age  of  7  years  the  weight  of  the 
brain  already  averages  40  oz.,  and  about  14  years  the  brain  not  infrequently 


WEIGHT  OF  THE  BRAIN  AND  CORD 


633 


m 


CZ 


i      n 


m 


Ix'-J 


IV 


Sb! 


.  .Tangential  fibers  (Vic 
d  Azyr's  ribbon) 


.  .Striae  of  Bechterew 


. .  Superradiary     network      (of 
the  second  and  third  layers) 


.  .Striae  of  Baillarger 


. .  Intermediary     network    (of 
the  third  and  fourth  layers) 


.  .Meynert's  intracortical  asso- 
ciation fibers 


. .  Subcortical  association  fibers 


FIG.  397.  Fio.  398. 

FIG.  397. — Schematic  Diagram  of  the  Different  Layers  of  the  Cerebral  Cortex.  (After 
Ramon  y  Cajal.)  I,  II,  III,  and  IV,  Layers  of  cortical  cells.  M,  Molecular  layer;  pPy, 
layer  of  small  pyramidal  cells;  gPy,  layer  of  large  pyramidal  cells;  Pm,  layer  of  poly- 
morphous cells. 

FIG.  398. — Schematic  Diagram  Showing  the  Arrangement  of  the  Nerve  Fibers  in  the 
Cerebral  Cortex.  The  dotted  lines  separate  the  four  cellular  layers  of  Cajal.  Sb,  White 
substance. 


634  THE    NERVOUS    SYSTEM 

reaches  the  weight  of  48  oz.  Beyond  the  age  of  forty  years  the  weight  slowly 
but  steadily  declines  at  the  rate  of  about  i  oz.  in  10  years. 

The  average  weight  of  the  female  brain  is  less  than  the  male ;  and  this  dif- 
ference persists  from  birth  throughout  life.  The  difference  amounts  to  about 
5  oz.  Thus  the  average  weight  of  an  adult  woman's  brain  is  about  44  oz. 

The  brains  of  idiots  are  generally  much  below  the  average,  some  weighing 
less  than  1 6  oz.  Still  the  facts  at  present  collected  do  not  warrant  more  than 
a  very  general  statement,  to  which  there  are  numerous  exceptions,  that  the 
brain  weight  corresponds  to  some  extent  with  the  degree  of  intelligence. 


Cfl 


FIG.  399. — Brain  of  the  Orang,  §  Natural  Size,  Showing  the  Arrangement  of  the 
Convolutions.  Sy,  Fissure  of  Sylvius;  R,  fissure  of  Rolando;  EP,  external  perpendicular 
fissure;  Olf,  olfactory  lobe;  Cb,  'cerebellum;  PV,  pons  Varolii;  MO,  medulla  oblongata. 
As  contrasted  with  the  human  brain,  the  frontal  lobe  is  short  and  small  relatively,  the  fissure 
of  Sylvius  is  oblique,  the  temporo-sphenoidal  lobe  very  prominent,  and  the  external  per- 
pendicular fissure  very  well  marked.  (Gratiolet.) 

There  can  be  little  doubt  that  the  complexity  and  depth  of  the  convolutions, 
which  indicate  the  area  of  the  gray  matter  of  the  cortex,  correspond  with  the 
degree  of  intelligence. 

The  spinal  cord  of  man  weighs  from  i  to  i  \  oz. ;  its  weight  relatively  to  the 
brain  is  about  1:40  in  the  adult.  As  we  descend  the  animal  scale,  this  ratio 
constantly  increases  till  in  the  mouse  it  is  1:4.  In  cold-blooded  animals 
the  relation  is  reversed,  the  spinal  cord  is  the  heavier.  In  the  newt,  1:105; 
and  in  the  lamprey,  j  :  133. 

The  most  distinctive  points  in  the  human  brain,  as  contrasted  with  that  of 
apes,  are:  i.  The  much  greater  size  and  weight  of  the  whole  brain.  The  brain 
of  a  full-grown  gorilla  weighs  only  about  15  oz.  (450  grms.),  which  is  less  than 
one-third  of  the  weight  of  the  human  adult  male  brain,  and  barely  exceeds 
that  of  the  human  infant  at  birth.  2.  The  much  greater  complexity  of  the 
convolutions,  especially  the  existence  in  the  human  brain  of  tertiary  convolu- 
tions in  the  sides  of  the  fissures.  3.  The  greater  relative  size  and  complexity 
and  the  blunted  quadrangular  contour  of  the  frontal  lobes  in  man,  which  are 
relatively  broader,  longer,  and  higher  than  in  apes.  In  apes  the  frontal  lobes 
project  keel-like  (rostrum)  between  the  olfactory  bulbs.  4.  The  much  greater 


GENERAL   FUNCTIONS    OF    THE    CEREBRUM  635 

prominence  of  the  temporo-sphenoidal  lobes  in  apes.  5.  The  fissure  of  Sylvius 
is  nearly  horizontal  in  man,  while  in  apes  it  slants  considerably  upward.  6. 
The  distinctness  of  the  fissure  of  Rolando. 

Most  of  the  above  points  are  shown  in  the  accompanying  figure  of  the  brain 
of  the  orang. 

GENERAL  FUNCTIONS  OF  THE  CEREBRUM. 

Evidence  regarding  the  physiology  of  the  cerebral  hemispheres  has 
been  obtained,  as  in  the  case  of  other  parts  of  the  nervous  system,  from  the 
study  of  anatomy,  from  pathology,  and  from  experiments  on  the  lower  ani- 
mals. The  chief  evidences  regarding  the  functions  of  the  cerebral  hemi- 
spheres derived  from  these  various  sources  are  briefly  these:  i.  Any  severe 
injury  of  them,  such  as  a  general  concussion,  or  sudden  pressure  as  by  apo- 
plexy, may  instantly  deprive  a  man  of  all  power  of  manifesting  externally 
any  mental  faculty.  2.  In  the  same  general  proportion  as  the  higher  mental 
faculties  are  developed  in  the  vertebrates  and  especially  in  man  at  different 
ages,  as  well  as  in  different  individuals,  the  greater  is  the  development  of 
the  cerebral  hemispheres  in  comparison  with  the  rest  of  the  cerebro-spinal 
system.  3.  No  other  part  of  the  nervous  system  bears  a  corresponding 
proportion  to  the  development  of  the  mental  faculties.  4.  Congenital  and 
other  morbid  defects  of  the  cerebral  hemisphere  are,  in  general,  accompanied 
by  corresponding  deficiency  in  the  range  or  power  of  the  intellectual  faculties 
and  the  higher  instincts.  5.  Removal  of  the  cerebral  hemispheres  in  the 
lower  animals  produces  effects  corresponding  with  what  might  be  antici- 
pated from  the  foregoing  facts. 

Effects  of  the  Removal  of  the  Cerebrum.-— The  removal  of  the  cere- 
brum in  the  lower  animals  appears  to  reduce  them  to  the  condition  of  a 
mechanism  without  spontaneity. 

In  the  case  of  the  frog,  when  the  cerebral  lobes  have  been  removed,  the  ani- 
mal appears  similarly  deprived  of  all  power  of  spontaneous  movement.  But 
it  sits  up  in  a  natural  attitude  and  breathes  quietly.  When  pricked  it  jumps 
away.  When  thrown  into  the  water  it  swims.  When  placed  upon  a  board 
it  remains  motionless,  although,  if  the  board  be  gradually  tilted  over  till  the 
frog  is  on  the  point  of  losing  his  balance,  he  will  crawl  up  till  he  regains  his 
equilibrium  and  comes  to  be  perched  quite  on  the  edge  of  the  board. 

If  the  frog  be  turned  on  his  back,  he  regains  his  normal  position.  If  his 
back  is  stroked  gently  he  will  utter  the  usual  croaking  sound.  These  activi- 
ties are  carried  on  by  the  normal  frog.  There  is  one  striking  difference, 
however,  between  the  brainless  frog  and  the  normal:  the  former,  if  placed 
in  a  position  and  left  undisturbed,  will  remain  without  moving  for  an 
indefinite  time.  It  has  apparently  lost  the  power  to  initiate  movements. 
Presumably  any  memory  impressions  or  effects  of  former  experiences  have 
been  lost.  Even  the  more  elemental  stimuli,  which  come  from  tissue 


636  THE    NERVOUS    SYSTEM 

hunger  and  thirst,  apparently  do  not  affect  the  brainless  frog.  In  other 
words,  the  operation  has  reduced  the  animal  to  the  condition  of  an  autom- 
aton capable  of  carrying  on  complex  activities,  but  only  upon  receiving 
some  definite  stimulus.  This  condition  contrasts  sharply  with  that  resulting 
from  the  removal  of  the  entire  brain,  leaving  only  the  spinal  cord.  In  this 
spinal  cord  frog  only  the  simpler  reflex  actions  can  take  place.  The  frog 
does  not  breathe.  It  lies  flat  on  the  table  instead  of  sitting  up.  When 
thrown  into  a  vessel  of  water  it  sinks  to  the  bottom.  When  its  legs  are 
pinched  it  kicks  out,  but  does  not  leap  away  as  in  the  normal. 

If  the  cerebrum  of  the  frog  be  removed,  taking  special  care  not  to  interfere 
with  the  optic  nerves  or  the  thalami,  then  it  acts  somewhat  differently. 
Whereas  with  the  entire  cerebrum  removed  it  makes  no  effort  to  take  food,  in 
this  instance  it  is  said  to  attempt  to  catch  flies  or  other  insects,  and  will  show 
other  signs  of  spontaneous  activity.  It  will  avoid  an  object  and  shows  signs  of 
responding  to  visual  sensations,  such  as  the  attempt  to  feed  just  mentioned. 

The  cerebral  lobes  of  the  frog,  however,  are  very  low  in  the  scale  of  de- 
velopment as  compared  with  other  vertebrates.  The  cortex  is  a  simple 
layer  of  rather  small  cells,  and  the  total  volume  of  the  cortex  as  compared 
with  other  portions  of  the  brain  is  small. 

The  case  of  the  pigeon,  which  represents  a  higher  animal  in  the  scale, 
has  been  extensively  studied  by  Flourens  and  others.  They  have  shown 
that  when  the  cerebrum  is  carefully  removed,  leaving  the  basal  nuclei  un- 
disturbed, and  the  animal  has  recovered  from  the  immediate  effects  of  the 
shock,  it  is  able  to  carry  on  many  co-ordinate  activities.  In  the  first  place  it 
can  stand  or  perch  without  difficulty.  If  placed  on  its  back  it  immediately 
regains  its  equilibrium.  If  tossed  in  the  air  it  flies  until  it  comes  in  contact 
with  a  firm  support.  If  disturbed  on  its  perch  it  will  walk  away,  showing 
the  power  to  co-ordinate  not  only  wing  muscles,  but  the  leg  muscles.  If  left 
undisturbed,  such  a  pigeon  will  occasionally  make  motions,  i.e.,  open  its 
eyes,  move  its  head,  preen  its  feathers,  or  even  take  a  step  or  two.  It 
spends  most  of  its  time,  however,  sitting  quietly  as  though  asleep.  If  aroused, 
the  animal  shows  little  or  no  signs  of  excitement  or  fright. 

After  several  months  such  pigeons  are  usually  said  to  increase  the  motions 
of  spontaneity  or  take  short  flights,  avoiding  obstacles  in  the  way  and  alight- 
ing definitely  on  the  perch.  They  will  pick  around  among  food  for  definite 
articles,  apparently  attempting  to  select  the  food.  Early  after  the  operation 
the  pigeon  will  pick  at  objects  indiscriminately,  but  does  not  take  food  un- 
less it  is  placed  in  the  mouth. 

Apparently  the  main  effect  produced  here  is  to  diminish  the  complexity 
and  efficiency  of  those  activities  which  we  call  spontaneous.  The  surprising 
thing  is  that  there  is  as  little  disturbance  among  the  motor  functions  as  is 
found. 

In  mammals  it  is  difficult  to  remove  the  cerebral  hemispheres,  but  in  those 


REACTIONS    OF    THE    HUMAN    WITHOUT    CEREBRUM  637 

animals  in  which  the  operation  has  been  carried  out,  in  the  rabbit  and  rat, 
a  result  very  similar  to  those  observed  in  the  case  of  the  frog  and  pigeon  has 
been  obtained.  The  animal  is  able  to  maintain  its  equilibrium,  to  run  or 
jump,  and  in  fact  successfully  carry  out  the  most  complicated  co-ordinated 
movements,  but  it  is  unable  to  originate  them  without  stimulation.  In  the 
case  of  the  dog,  it  has  been  found  impossible  to  remove  the  whole  brain  at  one 
operation.  However,  Goltz  has  succeeded  in  removing  both  the  cerebral 
hemispheres  of  the  dog  by  doing  the  operation  in  successive  stages  and  taking 
extraordinary  precautions  to  protect  his  animal  against  the  great  fall  of 
temperature  and  the  immediate  shock  of  the  operation.  He  kept  his  dog 
alive  for  some  eighteen  months  and  secured  a  complete  recovery  from  the 
series  of  operations.  Goltz's  dog  was  able  to  walk  about,  it  responded  to  a 
bright  light  by  closing  its  eyes,  and  could  be  aroused  by  a  sharp,  loud  sound. 
It  spent  its  time  lying  down  in  the  cage,  sleeping  rolled  up  dog-fashion. 
When  aroused  by  stimulation  of  the  skin,  it  would  move  away  from  the 
stimulating  object  and  would  sometimes  growl  and  snap  at  the  object. 
If  it  snapped  at  the  object  it  would  do  so  without  going  toward  it  or  making 
the  usual  effort  to  seize  the  object  which  we  are  accustomed  to  expect  of  a 
normal  vicious  dog.  This  dog  did  not  spontaneously  feed  itself,  but  had 
to  have  food  placed  in  its  mouth  before  it  would  swallow.  But  the  animal 
finally  learned  to  take  food,  as  in  the  case  of  the  pigeon.  This  animal  gave 
very  definite  responses  to  its  condition  of  nourishment;  it  slept  quietly  and 
was  peaceful  when  fully  fed,  but  was  restless  and  irritable  when  hungry. 

Goltz' s  dog  showed  complete  absence  of  those  activities  which  we  would 
call  psychic.  That  is  to  say,  it  showed  no  memory  signs,  it  was  unable 
to  learn  the  signal  for  feeding,  it  did  not  manifest  any  fondness  or  signs  of 
pleasure  at  the  presence  of  its  caretaker.  In  short,  there  was  a  complete 
loss  of  memory  and  intelligence,  and  the  animal,  although  performing  some 
activities,  was  in  fact  reduced  to  a  mere  automaton.  It  would  be  difficult  to 
imagine  a  more  crucial  experiment  to  elucidate  the  function  of  the  cerebral 
cortex. 

It  is  quite  evident  that  the  apparatus  for  carrying  out  co-ordinated  move- 
ments in  these  animals  is  not  localized  either  in  the  cerebrum  or  in  the  spinal 
cord.  It  must  therefore  be  connected  in  some  way  with  the  parts  of  the 
brain  below  the  cerebrum  and  above  the  cord.  There  is  no  reason  why  such 
an  arrangement  may  not  be  supposed  to  exist  in  the  human  brain,  although 
we  must  look  upon  the  cerebrum  as  the  originator  of  voluntary  movements. 

The  Reactions  of  the  Human  without  Cerebrum. — In  1913  Edinger 
and  Fischer  reported  a  most  interesting  case  of  a  child  who  without 
cerebrum  lived  to  the  age  of  three  and  three-fourths  years,  see  figure  400. 

The  cerebrum  in  this  child  consists  only  of  a  mass  of  cysts  without  nerve 
tissue.  The  optic  and  olfactory  nerves  were  also  lacking  in  nerve  elements, 
though  the  optic  chiasma  had  some  nerve  tissue.  The  cerebellum  did  not 


THE    NERVOUS    SYSTEM 

differ  from  the  normal.  The  midbrain  was  diminished  in  size.  The  cor- 
pora quadrigemina  had  fewer  nerve  cells  and  fibers  than  the  normal.  The 
corpus  striatum  was  normal  on  one  side,  but  on  the  other  lacked  portions  of 
the  putamen  and  the  nucleus  caudatus.  The  thalamic  ganglia  which  are 
well  developed  in  a  normal  child  were  absolutely  lacking  except  to  a  slight 
extent  in  the  hypothalamic  nuclei  where  a  few  fibers  coming  from  the  striatum 
were  present.  Of  the  cerebrum  proper  not  the  least  trace  remained,  either 
macroscopic  or  microscopic  except  for  the  connective  tissue  vascular  frame- 
work. 


FIG.  400. 

This  child  showed  no  physiological  development  from  birth  until  death. 
In  the  language  of  the  authors,  "  It  was  astonishing  how  much  less  this  child 
could  accomplish  than  the  dog  without  a  cerebrum."  While  the  operated 
dog  showed  the  activities  described  above,  the  child  remained  quiet  without 
conscious  control  over  its  limbs  and  body.  It  did  not  grasp  with  the  hands, 
showed  only  a  certain  motility  of  the  face,  the  lips  were  co-ordinated  with 
the  tongue  in  sucking,  but  otherwise  the  restlessness  and  motility  shown  by  the 


THE    MOTOR    FUNCTION    OF    THE    CEREBRAL   CORTEX  639 

dog  were  entirely  absent  in  the  child.  The  child  kept  up  a  perpetual  crying 
from  the  second  year  on,  but  the  crying  could  be  stilled  by  pressure,  especially 
about  the  head.  The  child  had  no  control  over  its  functions  and  showed  no 
consciousness  of  physical  discomfort.  "This  child  without  cerebrum  was 
even  less  active  than  a  decerebrate  fish  or  frog." 

In  comparing  the  reactions  of  this  child  with  those  of  the  decerebrate  dog, 
it  must  be  remembered  that  the  operated  dog  has  not  only  a  more  mature  cord 
and  medulla,  but  also  an  intact  and  well  developed  thalamus.  The  nuclei 
of  the  thalamus  may  quite  possibly  carry  on  extensive  nerve  co-ordinations 
which  were  of  course  lacking  to  the  child.  However,  the  child  possessed  the 
necessary  basic  nervous  mechanism  for  the  control  of  respiration,  circulation, 
digestion,  alimentary  motion,  etc.,  adequate  for  the  purely  physical  life. 
The  case  is  only  another  link  in  the  chain  of  evidence  which  more  firmly 
establishes  the  view  that  the  entire  psychic  life  is  inseparably  associated  with  ' 
the  normal  physiological  functioning  of  the  cerebral  cortex. 

LOCALIZATION  OF  THE  MOTOR  FUNCTION  OF  THE  CEREBRAL 

CORTEX. 

The  experiments  upon  the  brains  of  various  animals  by  means  of  electrical 
stimulation  have  demonstrated  that  there  are  definite  regions  of  the  cerebral 
cortex  the  stimulation  of  which  produces  definite  movements  of  co-ordinated 
groups  of  muscles  of  the  opposite  side  of  the  body.  Fritsch  and  Hitzig  were 
the  first  to  show  that  the  cerebral  cortex  responds  to  electrical  irritation. 
They  employed  a  weak  constant  current  in  their  experiments,  applying  a 
pair  of  fine  electrodes  not  more  than  one- twelfth  inch  apart  to  different  parts 
of  the  cerebral  cortex.  The  results  thus  obtained  have  been  confirmed  and 
extended  by  Ferrier,  Sherrington,  and  many  others,  stimulating  chiefly  with 
induction  currents. 

The  fundamental  phenomena  observed  in  all  these  cases  may  be  thus 
epitomized: 

i.  Excitation  of  the  same  spot  on  the  cortex  is  always  followed  by  the 
same  movement  in  the  same  animal.  2.  The  area  of  excitability  for  any 
given  movement  is  extremely  small,  and  admits  of  very  accurate  definition. 
3.  In  different  animals  excitations  of  anatomically  corresponding  spots  pro- 
duce contractions  in  similar  or  corresponding  muscles. 

The  various  definite  movements  resulting  from  the  electric  stimulation 
of  circumscribed  areas  of  the  cerebral  cortex  are  enumerated  in  the  descrip- 
tion of  the  accompanying  figures  of  the  dog's  and  monkey's  brains. 

In  the  case  of  the  dog  the  results  obtained  are  summed  up  as  follows  by 
Hitzig:  i.  One  portion,  anterior,  of  the  convexity  of  the  cerebrum  is  motor; 
another  portion,  posterior,  is  non-motor.  2.  Electric  stimulation  of  the 
motor  portion  produces  co-ordinated  muscular  contraction  on  the  opposite 
side  of  the  body.  3.  With  very  weak  currents,  the  contractions  produced 


640 


THE    NERVOUS    SYSTEM 


are  distinctly  limited  to  particular  groups  of  muscles;  with  stronger  currents 
the  stimulus  is  communicated  to  other  muscles  of  the  same  or  neighboring 
parts.  4.  The  portions  of  the  brain  intervening  between  these  motor  centers 
are  inexcitable. 


FIG.  401. — Brain  of  Dog,  A,  Viewed  from  Above  and  B  in  Profile.  F,  Frontal  fissure 
sometimes  termed  crucial  sulcus,  corresponding  to  the  fissure  of  Rolando  in  man;  S, 
fissure  of  Sylvius,  around  which  the  four  longitudinal  convolutions  are  concentrically 
arranged;  i,  flexion  of  head  on  the  neck,  in  the  median  line;  2,  flexion  of  head  on  the  neck, 
with  rotation  toward  the  side  of  the  stimulus;  3,  4,  flexion  and  extension  of  anterior  limb; 
5,  6,  flexion  and  extension  of  posterior  limb;  7,  8,  9,  contraction  of  orbicularis  oculi  and  the 
facial  muscles  in  general.  (Dalton.) 

Following  strong  stimulation  of  cortical  motor  centers  other  groups  of 
muscles  than  those  innervated  by  the  centers  stimulated  may  also  take  part  in 
the  contractions. 

According  to  the  further  researches  of  Schafer  and  Horsley,  electrical 


MOTOR   AREAS    OF   THE   HUMAN  BRAIN 


641 


stimulation  of  the  marginal  convolution  internally  at  the  parts  corresponding 
with  the  ascending  frontal  and  parietal  convolutions,  from  the  front  back- 
ward, produces  movements  of  the  arm,  of  the  trunk,  and  of  the  leg. 

A  good  deal  of  doubt  was  thrown  upon  the  experiments  of  Ferrier  by 
Goltz  and  other  observers,  from  the  results  of  excising  the  so-called  motor 
areas  of  the  dog's  brain.  It  was  found  that  the  part  might  be  sliced  away  or 
washed  away  with  a  stream  of  water,  but  that  no  permanent  paralysis  ensued. 

More  extensive  observations,  however,  have,  in  the  main,  confirmed 
Ferrier' s  original  statement,  at  any  rate  with  regard  to  the  monkey's  brain. 
Destruction  of  the  motor  areas  for  the  arm  produces  some  permanent  paraly- 
sis of  the  arm  of  the  opposite  side,  and  similarly, of  that  for  the  leg,  paralysis 
of  the  opposite  leg.  If  both  areas  are  destroyed,  permanent  hemiplegia 


ANUS   A  VAGINA 


OPENING 
OF  JAW 


VOCAL 
CORDS 

MASTICATION 


FIG.  402. — Scheme  Showing  the  Motor  Areas  of  the  Brain.     (Adapted  from  Griinbaum 
and  Sherrington  by  Cunningham.) 

ensues.  Paralysis  of  so  extensive  and  permanent  a  character  does  not,  how- 
ever, appear  the  rule  when  the  brain  of  a  dog  is  used  instead  of  that  of  the 
monkey.  It  is  suggested  that  in  the  animal  lower  in  the  scale  the  functions 
which  in  the  monkey  are  discharged  by  the  cortical  centers  may  be  subserved 
to  a  greater  extent  by  the  basal  ganglia. 

Motor  Areas  of  the  Human  Brain. — It  is  naturally  of  great  impor- 
tance to  discover  how  far  the  results  of  experiments  upon  the  dog  and  monkey 
hold  good  with  regard  to  the  human  brain.  Evidence  furnished  by  diseased 
conditions  is  not  wanting  to  support  the  general  idea  of  the  existence  of  cor- 
tical motor  centers  in  the  human  brain,  figure  402. 

So  far,  however,  it  has  been  possible  to  localize  motor  functions  only 
in  the  precentral  convolutions  and  the  walls  of  the  adjacent  sulci. 


642 


THE    NERVOUS    SYSTEM 


The  relative  position  of  the  centers  is  probably  much  the  same  as  in  the 
monkey's  brain,  those  for  the  leg  above,  those  of  the  arm,  face,  lips,  and 
tongue  from  above  downward.  Destruction  of  these  parts  causes  paralysis, 
corresponding  to  the  district  affected,  and  irritation  causes  contractions  of 
the  muscles  of  the  same  part.  Again,  a  number  of  cases  are  on  record  in 
which  aphasia,  or  the  loss  of  power  of  expressing  ideas  in  words,  has  been 
associated  with  disease  of  the  posterior  part  of  the  lower  or  third  frontal  con- 


CAUOSUM 


COftP;GEN:iNT. 
SUP  QUADV  BODY 


PHALON 


TEMPORO-PONTINE 
TRACT 


LOBE. 


FIG.  403. — Diagram  of  Certain  Connections  of  the  Frontal,  Temporal,   and  Occipital 
Lobes.     Founded  on  the  observations  of  Flechsig,  Ferrier,  and  Turner.     (Cunningham.) 


volution  on  the  left  side.  This  condition  is  usually  associated  with  motor 
paralysis  on  the  right  side  of  the  body,  right  hemiplegia. 

This  district  of  the  brain,  particularly  the  convolutions  bounding  the 
fissure  of  Rolando  anteriorly,  is  now  generally  known  as  the  motor  area. 
There  is  now  no  doubt  whatever  that  this  area  gives  origin  to  the  nerve  fibers 
which  proceed  to  the  spinal  cord,  and  are  there  represented  as  the  pyramidal 
tracts. 

This  is  the  reason  that  movements  are  produced  on  stimulation  of  the 


MOTOR    AREAS    OF   THE   HUMAN  BRAIN 


643 


white  matter  after  the  superficial  gray  matter  of  the  animal's  brain  has  been 
sliced  off. 

These  motor  fibers  are  those  which  arise  from  the  pyramidal  cells  of  the 
cortex.  From  the  motor  area  of  the  cortex  they  converge  to  the  internal  cap- 
sules, and  pass  down  to  the  crus.  In  the  internal  capsule  the  fibers  which 
pass  to  the  pyramidal  tracts  of  the  spinal  cord  occupy  that  part  known  as 
the  knee  (genu)  and  the  anterior  two-thirds  of  the  posterior  limbs  of  the  cap- 
sule, figure  404.  In  this  district  the  fibers  for  the  face,  arm,  and  leg  are  in 
this  relation:  those  for  the  face  and  tongue  are  just  at  the  knee,  and  below  or 
behind  them  come  first  the  fibers  for  the  arm  and  then  those  for  the  leg. 

The  more  accurately  known  arrangements  of  these  fibers  in  the  monkey's 
brain,  named  in  order,  from  above  down,  are  those  for  the  eye,  head,  tongue, 


FIG.  404. — Showing    the    geniculo-calcarine    optic    pathway.     (Meyer,    copied    from 

Gushing.) 

mouth,  shoulder,  elbow,  digits,  abdomen,  hip,  knee,  digits.  These  fibers 
come  for  the  most  part  from  the  portion  of  the  cortex  in  front  of  the  fissure  of 
Rolando,  chiefly  from  the  precentral  gyrus,  hence  called  the  Rolandic  area. 
Those  fibers,  passing  between  the  occipital  lobe  and  the  thalamus  and 
superior  corpora  quadrigemina,  are  concerned  with  vision,  and  are  called 
fibers  of  the  optic  radiation.  In  like  manner,  from  the  inferior  corpora 
quadrigemina  and  the  internal  geniculate  bodies,  fibers  which  make  up  the 
auditory  radiation  pass  to  the  auditory  center  in  the  superior  temporal  gyrus. 
The  term  motor  centers  is  applied  to  cortical  areas  which  are  concerned 
with  the  development  of  voluntary  motor  impulses.  It  must  not  be  assumed 
that  the  motor  cells  of  the  area  initiate  such  impulses.  These  centers  react 
only  in  response  to  afferent  nerve  impulses  which  flow  in  upon  them.  This 


644  THE    NERVOUS    SYSTEM 

fact  was  conclusively  proven  by  Mott  and  Sherrington  who  cut  the  sensory 
roots  of  the  brachial  plexus  and  found  that  the  monkey  so  operated  lost  the 
power  of  voluntary  use  of  the  muscles  on  the  side  operated.  The  cortical 
motor  apparatus  here  remains  intact  but  lacking  the  necessary  sensory  stim- 
ulation the  cortical  cells  are  unable  to  initiate  the  motor  impulses  required  for 
voluntary  action. 

It  has  already  been  shown  that  the  motor  fibers  of  the  internal  capsule  of 
one  side  cross  over  to  the  opposite  side  in  the  decussation  of  the  pyramids  in 
the  medulla.  This  decussation  is  not  quite  complete,  as  some  fibers  pass 
down  on  the  same  side  in  the  direct  pyramidal  tract.  A  small  portion  of 
these  direct  fibers  end  around  the  motor  neurones  of  the  same  side,  but  the 
great  majority  cross  to  the  opposite  side  in  the  anterior  commissure  at  some 
lower  level  of  the  cord.  It  follows  that  the  motor  areas  of  the  cortex  on  one 
side  control  the  muscular  movements  of  the  opposite  side  of  the  body,  but 
also  to  a  slight  extent  those  of  the  same  side.  As  a  matter  of  fact  disease  in 
the  region  of  the  fissure  of  Rolando  is  usually  accompanied  by  a  disturbance 
of  the  motor  function  on  the  opposite  side  of  the  body,  although  there  is 
some  slight  motor  disturbance  on  the  same  side. 

LOCALIZATION   OF   SENSORY   FUNCTION   IN   THE  CEREBRAL 

CORTEX. 

There  is  evidence  that  fibers  from  the  nerves  of  special  sense  are  specially 
connected  with  definite  and  distinct  parts  of  the  cerebral  cortex,  the  sensory 
areas. 

The  fibers  from  the  sensory  nerves,  we  have  found,  are  connected  with 
the  cerebral  cortex  by  chains  of  neurones.  These  sensory  paths,  although 
complex,  are  definite  and  distinct.  Their  cortical  connections  have  been 
mapped  out  with  considerable  definiteness. 

The  Body  Sensory  or  Somesthetic  Area. — The  demonstrated  motor 
function  around  the  pre-Rolandic  region  for  a  long  time  obscured  the  fact 
that  this  region,  especially  the  pre-  and  post-central  convolutions,  is  inti- 
mately connected  with  the  perception  of  general  body  sensations.  Physi- 
ological and  pathological  observations  supported  this  view,  and  recently 
Flechsig  has  much  strengthened  the  view  by  his  method  of  studying  the 
progressive  development  of  the  brain.  In  figure  405  we  produce  Flechsig's 
diagram  showing  the  body  sensory  area,  aptly  designated  by  Barker  the 
somesthetic  area.  The  borders  of  the  area  are  more  or  less  indefinite  and  less 
distinct  than  the  main  portion.  This  is  indicated  in  the  figure  by  the  lighter 
shading.  Lesions  of  this  area  in  the  cortex  lead  to  loss  of  sensibility  in  defi- 
nite regions  of  the  opposite  side  of  the  body.  Reaction  in  this  cortical  area 
evidently  forms  a  necessary  link  in  the  chain  of  nerve  impulses  initiated  by 
cutaneous  sensory  stimulation  and  leading  to  conscious  sensations. 


VISUAL    OR    OPTIC    CENTER 


645 


Visual  or  Optic  Center. — The  termination  of  the  optic  nerve  in  each 
eye,  the  retina,  to  the  structure  of  which  we  shall  return  when  treating  of  the 
eye,  is  so  arranged  that  when  we  look  at  an  object  with  both  eyes,  symmetrical 
parts  of  the  retinae  are  used.  For  example,  if  we  examine  an  object  to  the 
left  of  the  center  of  vision,  an  image  of  that  object  is  focused  upon  the  right 
half  of  both  retinae,  viz.,  upon  the  temporal  side  of  the  right  retina,  and  upon 
the  nasal  side  of  the  left  retina.  The  optic  nerve  fibers  of  these  symmetrical 
parts  of  the  retinae  are  gathered  together  behind  where  the  optic  nerves  de- 
cussate, viz.,  in  the  optic  chiasma.  The  fibers  which  come  from  the  right 
side  of  both  eyes  are  contained  in  the  optic  tract  of  the  same  side,  viz.,  the 
right,  those  from  the  right  eye  being  outside  of  the  others.  In  the  same  way 
the  left  optic  tract  contains  internally  fibers  from  the  left  side  of  the  right  eye 
and  externally  those  from  the  left  side  of  the  left  eye.  The  optic  tract  thus 
formed  then  passes  backward  and  terminates  in  three  distinct  nuclei,  viz., 


FIG.  405. — Diagrams  to  Show  Flechsig's  Sensory  and  Association  Areas  on  the  Surface  of  the 
Cerebral  Hemisphere.     (From  Cunningham,  after  Flechsig.) 

the  pulvinar  of  the  thalamus,  the  anterior  corpus  quadrigeminum,  and  the 
lateral  corpus  geniculatum.  These  nuclei  atrophy  if  the  eyes  are  removed 
from  an  adult  animal;  and  if  the  eyes  are  removed  from  a  newly-born 
animal,  they  do  not  fully  develop.  Through  the  superior  corpora  quadri- 
gemina  the  optic  tract  establishes  synapses  that  bring  it  into  relation  with  the 
nucleus  of  the  third  nerve,  and  which  form  the  basis  of  the  eye  reflexes  to 
light  stimulation. 

It  appears  that  some  of  the  fibers  of  the  optic  tract  pass  directly  into  the 
cerebral  cortex  without  joining  with  the  thalamus,  corpus  quadrigeminum, 
or  corpus  geniculatum. 

It  was  shown  above  that  the  fibers  of  the  cerebral  cortex,  known  as  the 
optic  radiation,  pass  from  the  occipital  region  to  the  three  nuclei  about 
which  we  are  speaking,  viz.,  into  the  pulvinar  of  the  thalamus,  the 
anterior  corpus  quadrigeminum,  and  lateral  corpus  geniculatum,  and 
it  is  known  that  when  the  occipital  cortex  is  removed,  these  three 


646  THE    NERVOUS    SYSTEM 

atrophy.  It  has  been  further  shown  that  in  a  newly-born  animal  the 
removal  of  such  a  region  is  followed  by  imperfect  development  of  the  parts 
in  question. 

If  one  optic  nerve  be  divided,  blindness  of  the  corresponding  eye  results; 
but  if  one  optic  tract  be  divided  there  is  a  half  blindness  in  both  eyes,  which  is 
called  hemianopsia,  or  hemiopia,  right  or  left,  according  as  the  right  or  left 
field  of  vision  is  cut  off.  It  is  also  evident  that  the  occipital  lobe,  figures  403, 
406,  and  particularly  the  cuneus,  is  concerned  as  a  visual  center.  Not  only 
is  the  occipital  lobe  connected  with  the  optic  nerves,  as  we  have  seen,  but 
the  removal  of  the  right  occipital  lobe  in  an  animal  (monkey)  is  followed  by 
bilateral  hemiopia  of  the  right  retinae.  Removal  of  the  left  occipital  lobe  is 
followed  by  left  bilateral  hemiopia.  Removal  of  both  occipital  lobes  is  fol- 
lowed by  total  blindness.  Clinical  observations  give  convincing  proof  that 
in  man  also  the  occipital  lobes  are  the  cortical  centers  of  vision.  Disturb- 
ance of  the  optic  lobes  or  tumors  involving  the  optic  tracts,  figure  404,  lead  to 
disturbance  and  blindness  in  the  retinal  field. 

Olfactory  Center  in  the  Cortex. — The  olfactory  nerve  differs  from 
the  other  cranial  nerves.  In  reality  it  is  a  representative  of  the  olfactory  lobes 
of  other  animals,  which  are  part  of  the  cerebrum.  The  olfactory  lobe  origi- 
nates as  an  offshoot  from  the  cerebral  vesicle,  the  front  part  of  which  is  de- 
veloped into  the  bulb  of  the  olfactory  nerve,  while  the  back  part  forms  its 
peduncle.  The  nerve,  the  cavity  of  which  in  man  is  filled  up  in  the  fully  de- 
veloped condition  with  neurogliar  substance,  lies  upon  the  cribriform  plate 
of  the  ethmoid  bone,  and  is  contained  in  a  groove  on  the  under  surface  of 
the  frontal  lobe.  On  examination  of  the  lobe  it  is  found  to  be  thus  made  up: 
Beneath  the  neurogliar  layer  is  a  layer  of  longitudinal  fibers  and  a  few 
nerve  cells,  next  to  this  is  a  layer  of  small  cells,  nuclear  layer,  fibers  from  the 
layer  of  nerve  fibers  passing  through  it. 

The  nuclear  layer  is  also  separated  into  groups  of  cells  by  an  interlacing 
of  the  fibers.  The  next  layer  is  thick  and  is  composed  of  neuroglia  and  nerve 
fibers,  some  of  which  are  medullated,  as  well  as  of  cells  more  or  less  pyramidal 
in  shape.  Below  this  layer  is  the  layer  of  olfactory  glomeruli.  These  glomer- 
uli  are  small  synapses  of  olfactory  fibers.  The  larger  also  includes  small 
cells  and  granular  matter.  A  further  description  of  the  anatomy  of  these 
parts  is  given  later,  page  656. 

Fibers  of  the  olfactory  nerve  proper  are  found  below  this  layer,  and  pass 
through  the  cribriform  plate  to  be  distributed  to  the  olfactory  mucous  mem- 
brane. They  arise  from  cells  in  the  olfactory  mucous  membrane,  and  end  in 
the  glomeruli.  The  peduncle  of  the  nerve  or  the  olfactory  tract,  as  it  is  some- 
times called,  is  made  up  of  longitudinal  fibers  originating  in  the  bulb,  with 
neuroglia  and  some  nerve  cells. 

The  fibers  of  the  olfactory  tract  have  been  traced  into  the  nucleus  amyg- 
dalae and  its  juncture  with  the  hippocampal  gyrus  in  the  temporal  lobe, 


AUDITORY    CENTER   IN    THE    CORTEX 


647 


figure  392.  The  hippocampus  must  be  in  some  way  connected  with  smell, 
since  a  lesion  of  it,  leaving  the  olfactory  tract  uninjured,  seriously  interferes 
with  that  sense. 

Taste  Center  of  the  Cortex. — It  is  very  uncertain  where  the  taste  center 
is  situated,  if  such  exist.  It  has  been  placed  in  the  anterior  portion  of  the 
inferior  temporal  convolution,  not  far  from  that  of  smell,  figure  405. 

Auditory  Center  in  the  Cortex. — This  center  has  been  localized  in 


FIG.  406. — Scheme  of  the  Central  Connections  of  the  Optic  Fibers.     (Cunningham.) 


the  superior  temporal  convolution.  Experiments  have  been  made  which 
connect  auditory  impulses  on  either  side  with  the  inferior  corpus  quadri- 
geminum  and  the  median  corpus  geniculatum,  for  when  the  internal  ear  is 
destroyed  there  results  atrophy  of  these  bodies  as  well  as  of  the  lateral  fillet 
of  the  opposite  side.  On  the  other  hand,  destruction  of  the  part  of  the  tem- 
poral lobe  above  indicated  is  similarly  followed  by  atrophy  of  the  nuclei  of 
the  same  side.  These  nuclei  bear  much  the  same  relation  to  the  sense  of 
hearing  as  do  the  superior  corpora  quadrigemina  and  the  lateral  corpora 
geniculata  to  the  sense  of  sight,  figures  406  and  414. 


648  THE    NERVOUS    SYSTEM 

ASSOCIATION  CENTERS  OF  THE  CEREBRAL  CORTEX. 

The  theory  of  localization  of  the  function  of  different  parts  of  the  cerebral 
cortex  has  received  substantial  support  from  the  study  of  the  motor  and  the 
sensory  areas  in  man  and  the  mammals.  But  when  the  exploration  of  the 
cortex  is  complete  and  the  motor  and  sensory  areas  are  bounded  as  definitely 
as  may  be,  there  still  remain  great  areas  in  which  stimulation  is  apparently 
non-effective  so  far  as  our  present  means  of  interpretation  reveal.  Trau- 
matic and  pathological  lesions  produce  no  sensory  or  motor  disturbances. 
The  areas  of  this  class  which  are  most  extensive,  i.e.,  which  cover  the  greatest 
amount  of  cortex,  are  three  in  number — the  frontal  lobe,  the  parietal  lobe, 
and  a  large  part  of  the  temporal  lobe  below  the  superior  temporal  convolution. 

Flechsig  has  made  a  study  of  the  development  of  the  human  brain,  paying 
especial  attention  to  the  progressive  development  of  the  great  tracts  of  fibers. 
He  has  shown  that  the  tracts  appear  in  a  certain  order  of  sequence,  also  that 
the  myelinization  takes  place  progressively. 

These  observations  give  justification  to  the  assumption  that  effective 
functionization  is  attained  with  the  acquirement  of  the  myelin  sheath.  He 
showed  a  close  correspondence  in  time  between  the  development  of  the 
tracts  and  the  manifestation  of  functions  known  to  involve  the  tracts  in 
question.  The  great  somesthetic  area  and  its  tracts  are  first  to  develop; 
then  tracts  to  the  occipital  or  visual  center,  to  the  auditory  and  other  sensory 
centers,  and,  finally,  to  those  great  areas  whose  functions  remain  obscure. 

Basing  his  deductions  on  the  facts  of  development,  on  the  isolated  cases  of 
lesion  and  disease,  and  on  the  careful  comparative  studies  of  the  brains  of 
certain  men  of  unusual  intellectual  powers,  whose  personal  characteristics 
and  intellectual  genius  are  known,  Flechsig  has  advanced  the  hypothesis 
that  the  areas  of  the  cortex  not  concerned  directly  with  motor  or  sensory 
functions  are  association  areas. 

The  Association  Centers  of  Flechsig. — The  great  association  centers 
are  the  frontal,  parietal,  and  temporal,  figure  405.  These  regions  of  the 
cortex  are  apparently  not  directly  connected  with  tracts  of  the  brain  stem 
and  cord,  but  they  are  richly  connected  with  the  areas  that  do  have  connec- 
tion with  the  cord.  Short  association  fibers  connect  neighboring  convolu- 
tions within  the  centers,  fibers  which  are  chiefly  the  axones  of  the  poly- 
morphous cells  of  the  fourth  layer  of  the  cortex.  Long  association  fibers 
run  from  one  center  to  another,  such  as  the  cingulum,  superior  and  inferior 
longitudinal  fasciculi,  etc.  The  longer  connectives  run  from  association 
to  association  centers,  and  from  association  to  sensory  and  motor  centers. 
Flechsig  believes  that  the  sensory  centers  are  not  connected  directly  with 
each  other,  but  only  indirectly  through  the  association  areas. 

Cases  of  injury  and  of  disease  of  the  human  brain  in  the  association  areas 
are  not  numerous,  but  such  as  there  are  tend  to  confirm  Flechsig's  hypothesis 
that  the  function  of  these  areas  is  that  of  the  higher  psychic  activity. 


THE   ANTERIOR    OR   FRONTAL   ASSOCIATION   CENTER  649 

The  Anterior  or  Frontal  Association  Center. — The  frontal  area  is 
more  closely  connected  with  the  motor  areas  and  the  centers  for  the  somes- 
thetic  sense.  With  injury  to  this  area  the  individual  shows  weakness  in  at- 
tention, in  reflection,  and  in  control  over  the  expressions  of  anger,  self- appre- 
ciation, and  other  activities  that  are  expressive  of  personal  volitions  and 
emotions. 

The  American  crowbar  case  is  a  classical  instance  of  lesion  of  the  frontal 
lobe.  A  young  man  of  twenty-five  had  an  iron  bar,  an  inch  and  a  quarter 
in  diameter  and  over  three  feet  long,  driven  through  his  skull  and  brain 
by  the  premature  explosion  of  a  blast  of  powder.  He  not  only  recovered, 
but  lived  for  twelve  years  afterward.  At  the  post-mortem  examination  the 
puncture  was  found  to  be  through  the  prefrontal  lobe,  anterior  to  the  coro- 
nal suture. 

This  man  was  considered  a  most  efficient  workman  and  foreman  before 


FIG.  407. — The  Association  Fibers  in  the  Centrum  Ovale.  A,  Between  adjacent  con- 
volutions; B,  between  frontal  and  occipital  lobes;  C,  between  frontal  and  temporal  lobes, 
the  cingulum;  D,  between  temporal  and  frontal  lobes — lesion  of  this  tract  causes  paraphasia; 
E,  between  occipital  and  temporal  lobes — lesion  of  this  tract  causes  word-blindness;  C.N, 
caudate  nucleus;  O.T,  optic  thalamus. 

the  injury.  After  his  recovery  he  was  fitful,  impatient  of  restraint,  capri- 
cious, obstinate.  He  was  most  inconsiderate  of  his  associates,  profane  and 
passionate.  From  a  shrewd  business  man  he  was  changed  to  the  intellectual 
level  of  a  child  and  was  regarded  by  his  associates  as  mentally  unbalanced. 

A  summary  of  fifty  cases  of  pathological  lesions  of  the  prefrontal  areas  of 
the  human  brain  is  given  by  Williamson.  The  mental  traits  of  thirty-two  are 
summarized  in  the  following  terms:  "A  condition  of  mental  decadence;  a 
dull  mental  state;  loss  of  power  of  attention;  loss  of  memory;  loss  of  spon- 
taneity; the  patient  takes  no  heed  of  his  surroundings;  sleeping  during  the 
greater  part  of  the  day,  or  remaining  semi-comatose."  Yet  these  patients 
are  able  to  walk  about  and  execute  well  co-ordinated  muscular  activities  of 
all  kinds  that  do  not  involve  complex  intellectual  activity. 


650  THE    NERVOUS    SYSTEM 

The  Parietal  Association  Center. — Special  mention  is  made  of  this 
association  area  because  there  is  increasing  evidence  that  it  is  the  parietal 
region  of  the  brain,  rather  than  the  frontal,  as  popularly  believed,  that  is 
most  intimately  concerned  with  the  higher  acts  and  powers  of  imagination, 
idealization,  and  reasoning.  It  is  the  region  through  which  the  individual 
maintains  his  interests  and  relations  with  the  external  world  as  against  his 
own  body.  The  parietal  association  center  is  more  closely  related  to  the 
visual,  auditory,  and  speech  centers  of  the  cortex.  The  great  musician 
Bach  had  an  exceptionally  well  developed  parietal  region. 

On  the  Cortical  Centers  in  General. — For  purposes  of  clearness  in 
presentation,  the  cortical  centers  have  been  discussed  one  by  one,  but  the 
reader  is  guarded  against  the  thought  that  their  activities  are  in  any  sense 
isolated.  A  motor  area  does  not  usually  act  in  the  absence  of  sensory  or  af- 
ferent stimulation  in  the  actual  living  body,  whether  it  may  do  so  on  occa- 
sion or  not.  Neither  do  sensory  impressions  arising  in  the  peripheral  sense 
organ  make  their  way  over  definite  tracts  to  the  brain  and  cortex  and  arouse 
sensations  alone.  Sensations  do  not  occur  independent  of  motor  activities 
on  the  one  hand,  and  of  intellectual  acts  through  the  association  centers  on 
the  other. 

The  association  centers  are  the  highest  co-ordinating  regions  of  the  nervous 
system.  They  are  to  the  sensory  and  motor  centers  what  these  latter  are  to 
the  reflex  centers  of  the  cord.  The  difference  is  one  of  degree  and  not  of 
kind.  Further,  the  association  centers  are  probably  set  into  activity  by  the 
complex  of  inflowing  or  afferent  impulses  in  just  the  same  sense  that  the 
spinal  reflex  centers  are  set  in  activity  by  sensory  or  afferent  stimuli;  the 
condition  is,  of  course,  a  thousand  times  more  complex. 

THE  CRANIAL  NERVES. 

The  cranial  nerves  consist  of  twelve  pairs;  they  appear  to  arise  (super- 
ficial origin)  from  the  base  of  the  brain  in  a  bilateral  series,  which  extends 
from  the  under  surface  of  the  anterior  part  of  the  cerebrum  to  the  lower  end 
of  the  medulla  oblongata.  Traced  into  the  substance  of  the  brain  and 
medulla,  the  roots  of  the  nerves  are  found  to  take  origin  from  various  masses 
of  gray  matter. 

The  roots  of  the  first  or  olfactory  and  of  the  second  or  optic  nerves  are 
discussed  elsewhere.  The  third  and  fourth  nerves  arise  from  gray  matter 
beneath  the  corpora  quadrigemina;  and  the  roots  of  origin  of  the  remainder 
of  the  cranial  nerves  can  be  traced  to  gray  matter  in  the  floor  of  the  fourth 
ventricle,  and  in  the  more  central  part  of  the  medulla,  around  its  central 
canal,  as  low  down  as  the  decussation  of  the  pyramids. 

According  to  their  several  functions  the  cranial  nerves  may  be  thus 
arranged : 


Nerves  of  common  sensation 


THE  THIRD  NERVE  OR  MOTOR  OCULI 

f  Olfactory,    optic,    auditory,    parts    of 

Nerves  of  special  sense •<    the   facial,  glosso-pharyngeal,  and   of 

(^  the  trigeminal. 

{The  greater  portion  of  the  trigeminal, 
and  part  of  the  facial. 

(  The  motor  oculi,   trochlearis,  lesser  di- 
Nerves  of  motion -<    vision    of    the   tri-geminal,   abducens, 

(^   hypoglossal,  and  spinal  accessory. 
Mixed  nerves Facial,  glosso-pharyngeal,  and  vagus. 

The  physiology  of  the  first,  second,  and  eighth  nerves  will  be  considered 
with  the  Organs  of  Special  Sense,  see  also  figure  416. 

The  Third  Nerve  or  Motor  Oculi. — Origin. — The  third  nerve  arises  in 


FIG.  408. — Section  through  Superior  Corpus  Quadrigemmum  and  Part  of  the  Thal- 
amus.  s,  Aqueduct  of  Sylvius;  gr,  gray  matter  of  the  aqueduct,  c.q.  s.,  superior  corpus 
quadrigeminum;  /,  stratum  lemnisci;  o,  stratum  opticum,  c,  stratum  cinereum;  Th,  pulvinar 
of  optic  thalamus;  c.g.e,  c.g.i,  lateral  and  median  corpora  geniculata;  br.s,  br.i,  superior  and 
inferior  brachia'/,  fillet;  p.l,  posterior  longitudinal  bundle;  r,  raphe;  ///,  third  nerve,  and 
n.III,  its  nucleus;  l.p.p,  posterior  perforated  space;  s.n,  substantia  nigra — above  this  is  the 
tegmentum  with  the  circular  area  of  the  red  nucleus;  cr,  crusta;  II,  optic  tract;  M,  medullary 
center  of  hemisphere;  n.c,  nucleus  caudatus;  st,  stria  terminalis.  (After  Quain,  from 
Meynert.) 

three  distinct  bands  of  fibers  from  the  gray  nuclei  surrounding  the  aqueduct 
of  Sylvius  near  the  middle  line,  but  ventral  to  the  canal,  figure  408.  The 
nucleus  of  origin  consists  of  large  multipolar  ganglion  cells,  and  extends  to 
the  back  part  of  the  third  ventricle  as  far  as  the  level  of  the  superior  cor- 
pora quadrigemina.  The  fibers  pass  from  their  origin  partly  through  the 
red  nucleus  to  their  superficial  origin  in  front  of  the  pons  at  the  median  side 
of  each  crus.  The  third  nerve  does  not  decussate. 

Function. — The  third  nerve  supplies  the  levator  palpebrae  superioris  mus- 
cle, and  all  of  the  muscles  of  the  eyeball,  except  the  superior  oblique,  to 
which  the  fourth  nerve  is  distributed,  and  the  rectus  externus  which  re- 
ceives the  sixth  nerve.  Through  the  medium  of  the  ophthalmic  or  lenticular 


652 


THE    NERVOUS    SYSTEM 


ganglion,  of  which  it  forms  what  is  called  the  short  root,  it  also  supplies 
motor  filaments  to  the  iris  and  ciliary  muscle.  The  fibers  which  subserve 
the  three  functions,  accommodation,  contraction  of  the  pupil,  and  nerve- 
supply  to  the  external  ocular  muscles,  arise  from  three  distinct  groups  of 
cells.  Optic  reflexes  involving  movements  of  the  eyeballs  are  mediated 
through  fibers  from  cells  of  the  superior  corpora  quadrigemina  (which  re- 
ceive fibers  from  the  optic  nerve).  These  fibers  from  the  corpora  quadri- 
gemina descend,  chiefly  through  the  posterior  longitudinal  bundle,  figure 


FIG.  409. — Fourth  Ventricle  with  the  Medulla  Oblongata  and  the  Corpora  Quad- 
rigemina. The  Roman  numbers  indicate  superficial  origins  of  the  cranial  nerves,  while  the 
other  numbers  indicate  their  deep  origins,  or  the  position  of  their  central  nuclei.  8,  8',  8", 
Auditory  nuclei  nerves;  t,  funiculus  teres;  A,  B,  corpora  quadrigemina;  c.g,  corpus  genic- 
ulatum;  p.c.  pedunculus  cerebri;  m.c.p,  middle  cerebellar  peduncle;  s.c.p,  superior  cere- 
bellar  peduncle;  i.c.p,  inferior  cerebellar  peduncle;  l.c,  locus  ceruleus;  e.t,  eminentia  teres; 
a.c,  ala  cinerea;  a.n,  accessory  nucleus;  o,  obex;  c,  clava;  f.c,  funiculus  cuneatus;  f.g,  funicu- 
lus gracilis. 

383,   to  the  nuclei  of  the  third,  fourth,  and  sixth  nerves,  thus  rendering 
possible  co-ordinated  reflex  movements  of  all  the  eye  muscles. 

When  the  third  nerve  is  stimulated  within  the  skull,  all  those  muscles  to 
which  it  is  distributed  are  contracted.  When  it  is  paralyzed  or  divided,  the 
following  effects  ensue:  i.  The  upper  eyelid  can  be  no  longer  raised  by  the 
levator  palpebrae,  but  droops,  ptosis,  and  remains  gently  closed  over  the  eye, 
under  the  unbalanced  influence  of  the  orbicularis  palpebrarum,  which  is  sup- 
plied by  the  facial  nerve.  2.  The  eye  is  turned  outward  and  downward. 


THE    FIFTH    NERVE,    OR    TRIGEMINAL 


653 


external  strabismus,  by  the  unbalanced  action  of  the  rectus  externus  and  supe- 
rior oblique,  to  which  the  sixth  nerve  is  appropriated;  and  hence,  from  the 
irregularity  of  the  axes  of  the  eyes,  double  sight,  diplopia,  is  often  experienced 
when  a  single  object  is  within  view  of  both  the  eyes.  3.  The  eye  cannot  be 
moved  upward,  downward,  or  inward.  4.  The  pupil  becomes  dilated, 
mydriasis.  5.  The  eye  cannot  accommodate  for  short  distances. 

The  Fourth  Nerve,  or  Trochlearis. — Origin. — The  fourth  nerve 
arises  from  a  nucleus  consisting  of  large  multipolar  ganglion  cells  situated 
ventral  to  the  aqueduct  of  Sylvius,  and  the  inferior  corpus  quadrigeminum. 


VJJl 


FIG.  410. — Section  Across  the  Pons,  About  the  Middle  of  the  Fourth  Ventricle,  py, 
Pyramidal  bundles;  po,  transverse  fibers  passing  polt  behind,  and  po2,  in  front  of  py;  rt 
raphe;  o.s,  superior  olive;  a.V,  bundles  of  ascending  root  of  V.  nerve  enclosed  in  a  pro- 
longation of  the  substance  of  Rolando;  VI,  the  sixth  nerve;  nVI,  its  nucleus;  VII,  facial 
nerve;  Vll.a,  intermediate  portion;  nVII,  its  nucleus;  VIII,  auditory  nerve;  nVIII, 
lateral  nucleus  of  the  auditory.  (After  Quain.) 

The  fibers  from  both  sides  sweep  dorsally  around  the  central  gray  matter,  and 
reach  the  valve  of  Vieussens,  where  they  decussate  in  the  mid-line  of  the  roof, 
then  pass  forward  along  the  lateral  aspect  of  the  crus.  The  nucleus  of  the 
fourth  nerve  on  either  side  is  connected  with  those  of  the  third  and  sixth 
nerves  and  with  the  optic  reflex  center  previously  described. 

Functions. — The  fourth  nerve  is  exclusively  motor,  and  supplies  only  the 
trochlearis  or  superior  oblique  muscle  of  the  eyeball. 

The  Fifth  Nerve,  or  Trigeminal. — Origin. — The  fifth  or  trigeminal 
nerve  resembles  the  spinal  nerves  in  that  it  has  two  roots;  namely,  the  larger 
or  sensory,  in  connection  with  which  is  the  Gasserian  ganglion,  and  the  small 
or  motor  root,  which  has  no  ganglion,  and  which  passes  under  the  ganglion 
of  the  sensory  root.  The  fibers  of  origin  of  the  fifth  nerve  come  from  the 
floor  of  the  fourth  ventricle.  The  motor  root  arises  to  the  inside  of  the  sen- 
sory about  the  middle  of  each  lateral  half  of  the  fourth  ventricle.  The 


654  THE    NERVOUS    SYSTEM 

sensory  fibers,  however,  can  be  traced  down  in  the  medulla  oblongata  as  far 
as  the  upper  part  of  the  cord.  The  motor  nucleus  stretches  forward  as  far 
as  the  superior  corpus  quadrigemirium,  giving  rise  to  a  bundle  of  long  fibers 
termed  the  descending  root.  The  sensory  nucleus  receives  a  tract  of  sensory 
fibers  from  the  trigeminus  extending  as  low  as  the  second  cervical  nerve, 
and  this  forms  a  tract  at  the  tip  of  the  posterior  cornu,  between  it  and  the 
restiform  body.  The  cells  of  origin  of  the  sensory  tract  are  in  the  Gas- 


FIG.  411. — General  Plan  of  the  Branches  of  the  Fifth.  X  $.  i,  Lesser  root  of  the 
fifth;  2,  greater  root  passing  forward  into  the  Gasserian  ganglion;  3,  placed  on  the  bone 
above  the  ophthalmic  nerve,  which  is  seen  dividing  into  the  supra-orbital,  lachrymal,  and 
nasal  branches,  the  latter  connected  with  the  ophthalmic  ganglion;  4,  placed  on  the  bone 
close  to  the  foramen  rotundum,  marks  the  superior  maxillary  division,  which  is  connected 
below  with  the  spheno-palatine  ganglion,  and  passes  forward  to  the  infra-orbital  foramen; 
5,  placed  on  the  bone  over  the  foramen  ovale,  marks  the  inferior  maxillary  nerve,  giving 
off  the  anterior  auricular  and  muscular  branches,  and  continued  by  the  inferior  dental 
to  the  lower  jaw,  and  by  the  gustatory  to  the  tongue;  a,  the  submaxillary  gland,  the  sub- 
maxillary  ganglion  placed  above  it  in  connection  with  the  gustatory  nerve;  6,  the  chorda 
tympani;  7,  the  facial  nerve  issuing  from  the  stylomastoid  foramen.  (Charles  Bell.) 

serian  ganglion.  The  nerve  appears  at  the  ventral  surface  of  the  pons 
near  its  front  edge,  at  some  distance  from  the  mid-line. 

Motor  Functions. — The  first  and  second  divisions  of  the  nerve,  which 
arise  wholly  from  the  larger  root,  are  purely  sensory.  The  third  division  is 
joined  by  the  motor  root  of  the  nerve  and  is  of  course  both  motor  and  sensory. 

Motor  branches  of  the  lesser  or  non-ganglionic  portion  of  the  fifth  supply 
the  muscles  of  mastication,  namely,  the  temporal,  masseter,  two  pterygoid, 
anterior  part  of  the  digastric  and  mylohyoid.  Filaments  are  also  said  to 


THE    FIFTH   NERVE,    OR    TRIGEMINAL  655 

supply  the  tensor  tympani  and  tensor  palati  (Kolliker).  The  motor  func- 
tion of  these  branches  is  proved  by  the  violent  contraction  of  all  the  muscles 
of  mastication  in  experimental  irritation  of  the  third  or  inferior  maxillary 
division  of  the  fifth  nerve;  by  paralysis  of  the  same  muscles  when  the  nerve 
is  divided  or  disorganized;  and  by  the  retention  of  the  power  of  these  muscles 
when  the  facial  nerve  is  paralyzed.  Whether  the  branch  of  the  fifth  nerve 
which  is  supplied  to  the  buccinator  muscle  is  entirely  sensory,  or  in  part 
motor  also,  must  remain  for  the  present  doubtful.  From  the  fact  that  this 
muscle,  besides  its  other  functions,  acts  in  concert  or  harmony  with  the 
muscles  of  mastication  in  keeping  the  food  between  the  teeth,  it  might  be 
supposed  from  analogy  that  it  would  have  a  motor  branch  from  the  same 
nerve  that  supplies  them.  However,  the  so-called  buccal  branch  of  the  fifth 
is,  in  the  main,  sensory. 

Sensory  Functions. — All  the  anterior  and  antero-lateral  parts  of  the  face 
and  head,  with  the  exception  of  the  skin  of  the  parotid  region,  acquire  com- 
mon sensibility  through  branches  of  the  ganglionic  division  of  the  fifth  nerve. 
The  muscles  of  the  face  and  lower  jaw  acquire  muscular  sensibility  through 
the  filaments  of  the  ganglionic  portion  of  the  fifth  nerve  distributed  to  them 
with  their  proper  motor  nerves. 

Through  its  ciliary  branches  and  the  branch  which  forms  the  long  root 
of  the  ciliary  or  ophthalmic  ganglion,  it  exercises  some  influence  on  the 
movements  of  the  iris.  When  the  trunk  of  the  ophthalmic  portion  is  divided, 
the  pupil  becomes,  according  to  Valentine,  contracted  in  men  and  rabbits,  and 
dilated  in  cats  and  dogs,  but  in  all  cases  becomes  immovable  even  under  all 
the  varieties  of  the  stimulus  of  light.  How  the  fifth  nerve  affects  the  iris  is 
unexplained;  it  has  been  suggested  the  influence  of  the  fifth  nerve  on  the 
movements  of  the  iris  may  be  ascribed  to  the  affection  of  vision  in  conse- 
quence of  the  disturbed  circulation  or  nutrition  in  the  retina. 

Trophic  Influence. — The  morbid  effects  which  division  of  the  fifth  nerve 
produces  in  the  organs  of  special  sense  make  it  probable  that  the  fifth  nerve 
exercises  some  special  or  trophic  influence  on  the  nutrition  of  all  these  organs, 
although  the  effects  may  in  part  be  due  to  the  loss  of  sensibility  which  is  the 
natural  protective  safeguard.  Thus,  after  such  division  and  within  a  period 
varying  from  twenty-four  hours  to  a  week,  the  cornea  begins  to  be  opaque 
and  later  it  grows  completely  white.  A  low  destructive  inflammatory  process 
ensues  in  the  conjunctiva,  sclerotic  coat,  and  in  the  interior  parts  of  the  eye. 
The  sense  of  smell  may  be  at  the  same  time  lost  or  gravely  impaired.  Com- 
monly, whenever  the  fifth  nerve  is  paralyzed,  the  tongue  loses  the  sense 
of  taste  in  its  anterior  and  lateral  parts,  and  according  to  Gowers  in  the 
posterior  part  as  well. 

In  Relation  to  Taste. — The  tactile  sensibility  of  the  tongue  is  due  to  the  lin- 
gual branch  of  the  fifth  nerve,  which  supplies  the  anterior  and  lateral  parts  of 
the  tongue.  The  sense  of  taste  in  the  lateral  and  anterior  portions  of  the  tongue 


656 


THE    NERVOUS    SYSTEM 


has  recently  been  traced  back  to  the  pars  intermedia  and  chorda  tympani 
of  the  seventh,  figures  412  and  413.  It  forms  also  one  chief  sensory  link 
in  the  nervous  circle  for  reflex  action  in  the  secretion  of  saliva.  But,  de- 
ferring this  question  until  the  glosso-pharyngeal  nerve  is  to  be  considered, 
it  may  be  observed  that  in  some  brief  time  after  complete  paralysis  or  division 
of  the  fifth  nerve,  the  power  of  all  the  organs  of  the  special  senses  may  be  im- 
paired. They  may  lose  not  merely  their  sensibility  to  common  impressions, 


M.G 


I.C. 


OG. 


Sty.  hy 


FIG.  412. — The  Seventh  Nerve  and  Its  Branches.  VII,  Facial  nerve;  P.I,  pars 
intermedia;  VIII,  auditory  nerve;  Ag.Fal,  aqueduct  of  Fallopius;  G.G,  geniculate  ganglion; 
E.S.P,  external  superficial  petrosal  nerve;  M.M,  middle  meningeal  artery;  G.S.P,  great 
superficial  petrosal  nerve;  G.P.D,  great  deep  petrosal  nerve;  I.C,  internal  carotid  artery; 
Vid,  Vidian  nerve;  M.G.,  Meckel's  ganglion;  Ty.Pl,  tympanic  plexus;  S.D.P,  small  deep 
petrosal  nerve;  G.Ph,  Glosso-pharyngeal  nerve;  Ty,  tympanic  branch;  S.S.P.,  small  super- 
ficial petrosal  nerve;  O.G.,  optic  ganglion;  Stap,  nerve  to  stapedius;  C.T,  chorda  tympani 
nerve;  Z,,  lingual  nerve;  A. Va,  communication  with  auricular  branch  of  vagus;  P. A, 
posterior  auricular  nerve.  Sty.hy,  nerve  to  stylo-hyoid;  Di,  nerve  to  digastric  (posterior 
belly);  T.F,  temporal -facial  division;  C.F,  cervico-facial  division;  T,  temporal;  M,  malar; 
I.O,  infra-orbital;  B,  buccal,  S.M,  supra-mandibular;  I.M,  infra-mandibular  branches. 
(Cunningham.) 

for  which  they  all  depend  directly  on  the  fifth  nerve,  but  also  their  sensibility 
to  the  special  stimuli  to  which  they  are  adapted. 

The  Sixth  Nerve,  the  Abducens. — Origin. — The  sixth  nerve  arises 
from  a  compact  oval  nucleus,  situated  somewhat  deeply  at  the  back  part  of 
the  pons  near  the  middle  of  the  floor  of  the  fourth  ventricle.  The  eminentia 
teres  marks  its  position.  It  contains  moderately  large  cells  with  large  nerve 
axis-cylinder  processes.  It  is  connected,  figure  375,  with  the  nuclei  of  the 
third,  fourth,  and  seventh  nerves,  and  with  reflex  centers  of  the  optic  tracts, 
as  previously  mentioned.  The  root  is  thin,  and  passes  ventrally  and  laterally 


THE    SEVENTH   NERVE,    OR   FACIAL  657 

through  the  reticular  formation,  to  the  surface,  which  it  reaches  at  the  lower 
edge  of  the  pons,  opposite  the  front  end  of  the  pyramid. 

Functions. — The  sixth  nerve  is  exclusively  motor,  and  supplies  only  the 
rectus  externus  muscle  of  the  eye.  The  muscle  is  paralyzed  when  the  nerve 
is  divided.  In  all  such  cases  of  paralysis  the  eye  squints  inward  and  cannot 
be  moved  outward. 

The  Seventh  Nerve,  or  Facial. — Origin. — The  facial  or  seventh  pair  of 
nerves  arises  from  the  floor  of  the  central  part  of  the  fourth  ventricle,  behind 
and  in  line  with  the  motor  nucleus  of  the  fifth,  to  the  outside  of  and  deeper 
down  than  the  nucleus  of  the  sixth.  The  nucleus  is  narrower  in  front  than 
behind,  and  consists  of  large  motor  cells  with  well-marked  axis-cylinder  proc- 
esses, which  are  gathered  up  at  the  dorsal  surface  of  the  nucleus  to  form  a 
root.  The  root  describes  a  loop  around  the  nucleus  of  the  sixth  nerve, 
running  forward  for  some  little  distance  dorsal  to  the  nucleus,  then  descend- 
ing vertically,  passing  to  the  outside  of  its  own  nucleus,  between  it  and  the 


pars    intermed -j 

r.  auric. 


i 
petros .  sup.  maj. 

L motor  ET. 


FIG.  413. — Dissection  of  the  Sensory  and  Motor  Divisions  of  the  Facial  in  a  ao-cm.  Embryo 

(Pig).     (Streeter.) 

descending  root  of  the  fifth  nerve.  It  emerges  at  the  lower  margin  of  the 
pons,  lateral  to  the  sixth  nerve,  opposite  the  front  edge  of  the  groove  between 
the  olivary  and  restiform  bodies.  The  second  or  sensory  root  is  smaller  and 
is  called  the  pars  intermedia,  figure  412.  It  is  the  portion  which  is  connected 
with  the  chorda  tympani  and  geniculate  ganglion,  figure  413.  The  pars 
intermedia  terminates  within  the  brain  in  the  fasciculus  solitarius  in  com- 
mon with  the  glosso-pharyngeal. 

Functions. — The  seventh  nerve  is  the  motor  nerve  of  all  the  muscles  of  the 
face,  including  the  platysma,  but  not  including  the  muscles  of  mastication. 
It  supplies  the  stapedius,  the  stylo-hyoid  and  the  posterior  part  of  the 
digastric.  Its  branches  also  supply  the  rudimentary  muscles  of  the  external 
ear. 

Fibers  from  the  chorda  tympani  are  distributed  to  the  submaxillary 
gland  and  produce  secretion  when  stimulated. 

When  the  facial  nerve  is  divided  or  in  any  other  way  paralyzed,  the  loss  of 

42 


658 


THE    NERVOUS    SYSTEM 


function  in  the  muscles  which  it  supplies  interferes  with  the  perfect  exercise 
of  the  organs  of  the  special  senses.  Thus,  in  paralysis  of  the  facial  nerve  the 
orbicularis  palpebrarum  being  powerless,  the  eye  remains  open  through  the 
unbalanced  action  of  the  levator  palpebrse.  The  conjunctiva  is  thus  contin- 
ually exposed  to  the  air  and  dust  and  is  liable  to  repeated  inflammation, 
which  may  end  in  thickening  and  opacity  of  the  cornea. 


CORPOflA  QUADRIGEMINA 


FIG.  414. — The  Nuclei  of  Origin  and  Central  Connections  of  the  Auditory  and  Vestibular 

Nerve.     (Cunningham.) 

The  sense  of  taste  may  be  weakened  or  wholly  lost  in  paralysis  of  the 
facial  nerve,  which  involves  the  chorda  tympani.  This  result,  which  has 
been  observed  in  many  instances  of  disease  of  the  facial  nerve  in  man,  appears 
explicable  on  the  supposition  that  the  chorda  tympani  is  the  nerve  of  taste 
to  the  anterior  two-thirds  of  the  tongue,  its  fibers  being  distributed  with  the 
so-called  gustatory  or  lingual  branch  of  the  fifth.  Streeter  has  just  published 
a  study  of  the  development  of  the  seventh  and  eighth  nerves  in  which  he 
traces  the  chorda  tympani  through  the  pars  intermedia,  as  shown  in  figure 
413,  thus  settling  this  oft-disputed  question. 


THE    NINTH   NERVE,    OR    GLOSSO-PHARYNGEAL  659 

The  Eighth  Nerve,  or  Auditory. — The  eighth  nerve  consists  of  two 
divisions,  anatomically  distinct  and  functionally  independent.  These  are 
the  vestibular  and  the  cochlear  divisions  of  the  auditory  nerve. 

The  cochlear  division  arises  in  the  spiral  ganglion  and  passes  to  the 
medulla  to  establish  immediate  connections  with  the  ventral  cochlear  nucleus 
and  the  tuberculum  acusticum.  The  central  relations  of  these  nuclei  are 
established  by  the  striae  acusticae,  the  trapezoideus,  and  the  lateral  fillet  with 
the  internal  corpus  geniculatum  and  the  inferior  corpus  quadrigeminum  of 
the  opposite  side,  as  told  by  figure  414.  These  latter  nuclei  send  tracts  to 
the  auditory  center  in  the  superior  temporal  gyrus. 

The  vestibular  division  arises  in  the  vestibular  ganglion,  which  is  entirely 
distinct  from  the  cochlear  ganglion,  and  enters  the  medulla,  passing  to  the 
lateral  or  chief  auditory  nucleus.  From  this  point  the  relations  are  not  fully 
established,  but  apparently  fibers  pass  to  the  nucleus  fastigii  of  the  opposite 
side  and  to  the  vermis,  where  they  are  brought  into  relations  with  motor 
descending  paths. 

Functions. — The  cochlear  branch  is  the  auditory  nerve  proper,  and  the 
vestibular  is  the  nerve  of  equilibrium.  The  reader  is  referred  to  the  chapter 
on  Hearing  for  the  details  of  function. 

The  Ninth  Nerve,  or  Glosso-pharyngeal. — Origin. — The  glosso-phar- 
yngeal  nerves,  figure  378,  IX,  arise  by  nuclei  intimately  associated  with 
those  of  the  vagus  and  spinal  accessory  nerves.  The  union  of  the  nuclei  is 
indeed  so  intimate  that  it  will  be  as  well  to  consider  the  origins  of  the  ninth, 
tenth,  and  eleventh  nerves  together. 

These  three  nerves  emerge  from  the  bulb  and  spinal  cord  in  their  numerical 
order  from  above  downward,  the  bulbar  portion  from  the  lateral  aspect  of 
the  bulb  in  a  line  between  the  olivary  and  restiform  bodies;  and  the  spinal 
portion  from  a  line  intermediate  between  the  anterior  and  posterior  nerve 
roots  as  far  down  as  the  sixth  or  seventh  cervical  spinal  nerves. 

The  combined  glosso-pharyngeal-accessory-vagus  nucleus  appears  to  con- 
sist of  two  parts,  viz.,  one  median  or  common  origin,  having  conspicuous 
nerve  cells  of  moderate  size,  and  three  lateral  origins,  having  but  few  cells  of 
small  size.  These  are:  i,  the  nucleus  ambiguus,  which  lies  on  the  lateral  side 
of  the  reticular  formation  and  is  the  motor  origin  of  the  glosso-pharyngeal, 
the  vagus,  and  the  spinal  accessory;  2,  the  fasciculus  solitarius,  situated  in 
the  bulb,  ventral  and  a  little  lateral  to  the  combined  nucleus,  is  also  called  the 
ascending  root  of  the  glosso-pharyngeal  nerve  or  the  respiratory  bundle;  and 
3,  the  spinal  portion,  which  takes  origin  from  a  group  of  cells  lying  in  the  ex- 
treme lateral  margin  of  the  anterior  cornu.  This  is  the  origin  of  the  spinal 
accessory;  it  corresponds  to  the  antero-lateral  nucleus  of  the  bulb,  and  the 
lateral  part  of  the  gray  matter  of  the  spinal  cord. 

The  fibers  of  the  spinal  origin  of  the  nerve  pass  from  these  cells  through 
the  lateral  column  to  the  surface  of  the  cord.  The  fibers  from  the  median 


660  THE    NERVOUS    SYSTEM 

part  pass  in  a  ventral  and  lateral  direction  through  the  reticular  formation, 
then  ventral  to  or  through  the  gelatinous  substance  and  strand  of  fibers 
connected  with  the  fifth  nerve,  to  the  surface  of  the  bulb. 

The  fibers  from  the  nucleus  ambiguus  join  the  combined  nerve,  chiefly 
the  vagus  and  the  glosso-pharyngeal. 

The  bundles  of  fibers  of  the  fasciculus  solitarius  start  in  the  lateral  gray 
matter  of  the  cervical  cord  and  higher  in  the  reticular  formation  of  the  bulb, 
run  longitudinally  forward,  to  pass  into  the  roots  of  the  ninth  nerve.  It  is 
composed  of  sensory  fibers,  chiefly  of  the  glosso-pharyngeal,  and  of  the 
pars  intermedia  of  the  facial. 

The  glosso-pharyngeal  nerve  gives  filaments  through  its  tympanic  branch 
(Jacobson's  nerve),  to  the  fenestra  ovalis  and  fenestra  rotunda,  and  the  Eu- 
stachian  tube;  also  to  the  carotid  plexus,  and  through  the  petrosal  nerve,  to 
the  spheno-palatine  ganglion.  After  communicating  with  the  vagus  and, 
soon  after  it  leaves  the  cranium,  with  the  sympathetic,  with  the  digastric 
branch  of  the  facial,  and  the  accessory  nerve,  the  glosso-pharyngeal  divides 
into  the  two  principal  divisions  indicated  by  its  name.  These  divisions 
supply  the  mucous  membrane  of  the  posterior  and  lateral  walls  of  the 
upper  part  of  the  pharynx,  the  Eustachian  tube,  the  arches  of  the  palate, 
the  tonsils  and  their  mucous  membrane,  and  the  tongue  as  far  forward 
as  the  foramen  cecum  in  the  middle  line,  and  to  near  the  tip  at  the  sides  and 
inferior  part. 

Functions. — The  glosso-pharyngeal  nerve  contains  some  motor  fibers, 
together  with  fibers  of  common  sensation  and  the  sense  of  taste. 

Motor  fibers  are  distributed  to  the  palato-pharyngeus,  the  stylo-pharyn- 
geus,  palato-glossus,  and  constrictors  of  the  pharynx. 

Sensory  fibers  of  touch  and  of  common  sensation  are  distributed  to  the 
pharynx,  the  tonsils,  and  posterior  palate.  Nerves  of  taste  are  supplied  to 
the  taste  buds  on  the  posterior  third  of  the  tongue  and  to  the  fauces. 

The  Tenth  Nerve,  Vagus  or  Pneumogastric  Nerve. — The  origin  of  the 
vagus  nerve  is,  as  we  have  just  seen,  situated  in  the  lower  half  of  the  floor  of 
the  fourth  ventricle,  figure  378.  Its  nucleus  is  said  to  represent  the  homo- 
logue  of  the  cells  of  Clarke's  column  of  the  spinal  cord.  In  origin  it  is 
closely  connected  with  the  ninth,  eleventh,  and  the  twelfth.  The  com- 
bined glosso-pharyngeal- vago-accessory  nuclei  lie  outside  of,  close  to,  and 
parallel  with  the  nucleus  of  the  twelfth.  There  are  two  main  vagal  nuclei, 
one  motor,  the  other  sensory.  The  motor  nucleus  gives  rise  to  a  large 
group  of  efferent  fibers  of  wide  distribution  in  the  body  all  belonging  to  the 
bulbar  autonomies  of  Langley. 

Distribution. — It  has,  of  all  the  nerves,  the  most  varied  distribution  and 
functions,  either  through  its  filaments,  or  through  sensory  fibers  which 
are  mingled  in  its  branches,  and  give  rise  to  most  varied  reflex  reactions. 
The  vagus  supplies  sensory  branches,  which  accompany  the  sympathetic 


THE  TENTH  NERVE,  VAGUS  OR  PNEUMOGASTRIC  NERVE 


661 


FIG.  415. 


FIG.  416. 


FIG.  415. — The  Distribution  of  the  Tenth  or  Vagus  Nerve.  Va.R,  Va.L.,  Right  and 
left  vagi;  r,  ganglion  of  the  root  and  connections  with  Sy.,  sympathetic,  superior  cervical 
ganglion;  g.Ph.,  glosso-pharyngeal;  Ace.,  spinal  accessory  nerve;  m,  meningeal  branch;  Aur. 
auricular  branch;  t,  ganglion  of  the  trunk  and  connections  with  Hy.,  hypoglossal  nerve; 
Ci,  €2,  loop  between  the  first  two  cervical  nerves — Sy.,  sympathetic,  Ace.,  spinal  accessory 
nerve;  Ph.,  pharyngeal  branch;  Ph.PL,  pharyngeal  plexus;  S.L.,  superior  laryngeal  nerve; 
I.L.,  internal  laryngeal  branch;  E.L.,  external  laryngeal  branch;  I.C.,  internal,  and  E.G., 
external  carotid  arteries;  Cai,  superior  cervical  cardiac  branch;  Ca2,  inferior  cervical 
cardiac  branch;  R.L.,  recurrent  laryngeal  nerve;  Ca$,  cardiac  branches  of  recurrent  laryn- 
geal nerve;  Ca$,  thoracic  cardiac  branch  (right  vagus);  A. P. PL,  anterior,  and  P. P. PI., 
posterior  pulmonary  plexuses;  Oes.PL,  esophageal  plexus;  Gast.R.  and  Gast.L.,  gastric 
branches  of  vagus  (right  and  left);  Coe.Pl.,  celiac  plexus;  Hep.Pl.,  hepatic  plexus;  PI., 
splenic  plexus;  Ren.Pl ,  renal  plexus.  (Cunningham.) 

FIG.  416. — The  Constitution  of  the  Cardiac  Plexus.  Sy.,  Cervical  sympathetic  cord; 
C.i,  superior,  C.2,  middle,  and  C.3,  inferior  cervical  ganglia;  Car.i,  superior,  Car.2,  middle, 
and  Car.3,  inferior  cervical  cardiac  sympathetic  branches;  Va.,  vagus  nerve;  R.L.,  recurrent 
laryngeal  nerve;  s,  superior,  and  i,  inferior  cervical  cardiac  branches  of  vagus;  D.C.P., 
deep  cardiac  plexus;  S.C.P.,  superficial  cardiac  plexus;  A. P. P.,  anterior  pulmonary 
plexus;  P.P.P.,  posterior  pulmonary  plexus;  R.Car.P.,  right,  and  L.Car.P.t  left  coronary 
plexuses;  Art.Pul.,  pulmonary  artery.  (Cunningham.) 


662 


THE    NERVOUS    SYSTEM 


on  the  middle  meningeal  artery  supplying  the  back  part  of  the  meatus  and 
the  adjoining  part  of  the  external  ear.  It  is  connected  with  the  petrous 
ganglion  of  the  glosso-pharyngeal  and  with  the  spinal  accessory  which  con- 
tributes motor  fibers  for  the  larger  and  upper  portion  of  the  esophagus, 
and  inhibitory  fibers  for  the  heart.  It  is  connected  also  with  the  twelfth; 
with  the  superior  cervical  ganglion  of  the  sympathetic;  and  with  the  cer- 
vical plexus.  The  organs  supplied  by  the  branches  of  the  vagus  are  as 
follows : 

1.  A  large  portion  of  the  mucous  membrane  and  some  of  the  muscles 
of  the  pharynx  are  supplied  by  its  pharyngeal  branches,  through  the 
pharyngeal  plexus. 

2.  The  mucous  membrane  of  the  under  surface  of  the  epiglottis,  and  of 
the  greater  part  of  the  larynx,  and  the  crico-thyroid  muscle,  by  the  superior 
laryngeal  nerve. 

3.  The  mucous  membrane  and  muscular  fibers  of  the  trachea,  the  lower 
part  of  the  pharynx  and  larynx,  and  all  the  muscles  of  the  larynx  except  the 
crico-thyroid  are  supplied  by  the  inferior  laryngeal  nerve.     It  also  supplies 
the  first  segment  of  the  esophagus. 

4.  The  mucous  membrane  and  muscular  coats  of  the  esophagus  re- 
ceive fibers  from  the  esophageal  branches. 

5.  The  branches  of  the  vagus  form  the  supply  of  inhibitory  nerves  to 
the  heart  and  the  sensory  distribution  of  depressor  fibers  to  the  great 
arteries. 

6.  The  sensory  fibers  of  the  lungs  and  the  bronchial  muscle  motor 
nerves  are  supplied  through  the  anterior  and  posterior  pulmonary  plexuses. 

7.  The  stomach,  the  intestines,  the  spleen,  and  the  liver  are  supplied  by 
the  gastric,  splenic,  and  hepatic  vagus  branches. 

Functions. — Throughout  its  whole  course  the  vagus  contains  both  sen- 
sory and  motor  fibers.  To  summarize  the  many  functions  of  this  nerve, 
which  have  been  for  the  most  part  considered  in  the  preceding  chapters, 
it  may  be  said  that  it  supplies,  i,  motor  fibers  to  the  pharynx  and  esopha- 
gus, to  the  stomach  and  intestines,  to  the  larynx,  trachea,  bronchi,  and 
lungs;  2,  sensory  fibers  to  the  same  regions;  3,  inhibitory  fibers  to  the  heart; 
4,  inhibitory  afferent  fibers,  i.e.,  depressors,  to  the  vaso-motor  center. 

Surgical  division  of  both  vagi  or  of  both  their  recurrent  branches  is 
often  ultimately  fatal  in  young  animals;  but  in  old  animals  the  division  of 
the  recurrent  nerve  is  not  generally  fatal,  and  that  of  both  the  vagi,  even, 
is  not  always  fatal. 

The  Eleventh  Nerve,  or  Spinal  Accessory. — This  nerve  arises  by  two 
nuclei,  one  the  nucleus  ambiguus  from  a  center  in  the  floor  of  the  fourth 
ventricle,  partly  but  chiefly  in  the  medulla  and  continuous  with  the  glosso- 
pharyngeal- vagus  nucleus;  the  other,  from  the  outer  side  of  the  anterior 
cornu  of  the  spinal  cord  as  low  down  as  the  fifth  or  sixth  cervical  nerve. 


THE    AUTONOMIC    SYSTEM  663 

The  fibers  from  the  two  origins  come  together  at  the  jugular  foramen,  but 
separate  again  into  two  branches.  The  inner  arises  from  the  medulla  and 
joins  the  vagus,  to  which  it  supplies  fibers,  consisting  of  small  medullated 
nerve  fibers.  The  outer  consists  of  large  medullated  fibers  and  supplies  the 
trapezius  and  sterno-mastoid  muscles.  The  muscles  of  the  larynx,  all  of 
which  are  supplied,  apparently,  by  branches  of  the  vagus,  are  said  to  derive 
their  motor  nerves  from  the  accessory;  and  Vrolik  makes  the  very  signifi- 
cant statement  that  in  the  chimpanzee  the  internal  branch  of  the  accessory 
does  not  join  the  vagus  at  all,  but  goes  direct  to  the  larynx. 

The  Twelfth  Nerve,  or  Hypoglossal. — Origin  and  Connections. — The 
nerve  arises  from  a  large-celled  and  very  long  nucleus  in  the  bulb,  extending 
from  the  floor  of  the  fourth  ventricle  to  the  level  of  the  olivary  bodies  close 
to  the  mid-line  and  inside  the  nucleus  ambiguus.  Fibers  from  this  nucleus 
run  from  the  ventral  surface  through  the  reticular  formation  in  a  series  of 
bundles  passing  between  the  olivary  nucleus  laterally  and  the  pyramid  and 
accessory  olive  medially,  to  gain  the  ventral  surface.  The  nerve  emerges 
from  a  groove  between  the  pyramid  and  olivary  body.  The  fibers  of 
origin  are  continuous  with  the  anterior  roots  of  the  spinal  nerves. 

This  nerve  is  the  motor  nerve  to  the  muscles  connected  with  the  hyoid 
bone,  including  those  of  the  tongue.  It  supplies  the  sterno-hyoid,  sterno- 
thyroid,  and  omo-hyoid  through  its  descending  branch,  descendens  hypo- 
glossi;  the  thyro-hyoid  through  a  special  branch;  and  the  genio-hyoid, 
stylo-glossus,  hyo-glossus,  and  genio-hyoglossus  and  linguales  through  its 
lingual  branches. 

Functions. — The  function  of  the  hypoglossal  is  exclusively  motor.  In 
cases  of  hemiplegia  involving  the  functions  of  the  hypoglossal  nerve  the 
tongue  when  protruded  deviates  toward  the  paralyzed  side,  when  with- 
drawn it  turns  away  from  the  paralyzed  side.  Occasionally  it  is  not  pos- 
sible to  observe  any  deviation  in  the  direction  of  the  protruded  tongue; 
probably  because  the  tongue  is  so  compact  and  firm  that  the  muscles  on 
either  side  can  push  it  straight  forward  or  turn  it  for  some  distance  toward 
either  side.  In  hypoglossal  paralysis  from  cerebral  lesions  or  lesions  of  the 
peduncles  the  paralysis  is  contralateral. 

THE  AUTONOMIC  SYSTEM. 

In  the  introductory  statement  of  this  chapter  we  emphasized  the  fact 
that  the  outlying  anatomical  nerve  divisions  long  known  as  the  sympa- 
thetic system  is  an  organic  and  constituent  part  of  the  general  nervous 
system.  Histologically  there  is  close  relation  of  the  constituent  parts, 
embryonically  there  is  a  common  origin  of  the  neurones,  and  physio- 
logically the  mechanisms  are  dependency  related.  The  main  portion  of 
this  group  of  neurones  we  discuss  under  the  title  The  Autonomic  System. 


664 


THE   NERVOUS    SYSTEM 


In  the  discussion  the  intimate  anatomical  relationships  must  be  constantly 
kept  in  view. 

Organization  and  Distribution. — Strictly  speaking,  the  name  auto- 
nomic  nervous  system  emphasizes  physiological  groupings  of  nerve  cells 
and  pathways  rather  than  anatomical  structures.  However,  the  fact  is 
that  outside  the  brain  and  spinal  cord  and  exclusive  of  the  somatic  nerves 
there  are  numerous  nerve  ganglia.  These  ganglia  are  connected  by 
tracts  with  each  other  and  with  the  cerebro-spinal  axis. 


GrayRamus 
White  Ramus 

Sympathetic  Ganglion. 


ogiel 

mnathctic 

SOciatioa 

fiber 


Recurrent  Branc 

ofMeninges 


Sympalhetic  Ganglion. 


FIG.  417. — Schematic  Representation  of  the  Relation  of  the  Constituents  of  the  Sympathetic 
Chain  and  the  Spinal  Nerve.     (Modified  from  Hardesty,  in  Morris'  Anatomy.) 


The  chief  gross  constituent  unit  to  be  mentioned  is  a  bilaterally 
symmetrical  double  chain  of  ganglia  and  connecting  nerves  extending 
from  the  cranium  to  the  pelvis  and  along  each  side  of  and  in  front  of  the 
vertebral  column.  A  ganglion  is  connected  with  each  of  the  spinal  nerves 


GROSS    ANATOMICAL    RELATIONS  665 

below  the  region  of  the  first  dorsal,  but  there  are  only  three  cervical  ganglia 
in  the  neck  and  certain  ganglia  of  questionable  horn ology  in  the  head  region. 
This  chain  of  ganglia  is  known  in  the  historical  anatomical  literature  and 
in  the  older  physiological  literature  as  the  sympathetic  system. 

There  are,  however,  numerous  other  outlying  ganglia  such  as  the  gan- 
glia of  the  celiac  axis,  the  superior  mesenteric  and  hypogastric  plexuses,  as 
well  as  such  secondary  or  collateral  plexuses  as  the  aortic,  renal,  etc. 
From  these  ganglia  nerves  are  distributed  to  other  parts  of  the  system  and 
to  the  motor  end  mechanisms.  The  small  ganglia  in  connection  with 
those  branches  of  the  fifth  and  other  cerebral  nerves  which  are  distributed 
in  the  neighborhood  of  the  organs  of  special  sense,  namely,  the  ophthalmic, 
otic,  spheno- palatine,  and  submaxillary  ganglia,  etc.,  all  belong  to  the  cranial 
autonomies  as  we  shall  presently  see.  Similar  small  ganglia  are  found  on 
the  sacral  autonomic  pathways. 

Physiologically  there  are  two  functional  types  of  nerve  fibers,  afferent 
and  efferent,  which  run  in  autonomic  pathways.  The  former  or  afferent 
fibers  arise  in  the  sensory  epithelium  of  the  alimentary  canal,  lungs  or 
blood  vessels,  and  from  special  sense  organs  such  as  the  pacinian  corpuscles 
of  the  mesentery.  The  second  or  efferent  group  constitutes  the  autonomic 
system  proper.  It  is  to  this  group  that  the  cells  of  the  sympathetic  ganglia 
of  all  kinds  belong. 

There  is  still  a  third  order  of  nerve-cell  collections  or  ganglia  found  in 
the  walls  of  hollow  organs.  The  chief  and  best  known  of  these  ganglionic 
networks  are  found  in  the  walls  of  the  stomach  and  intestinal  canal  and  are 
known  as  the  plexuses  of  Auerbach,  and  of  Meissner.  Smaller  collections 
of  this  type  are  found  in  the  walls  of  the  urinary  bladder,  the  ureter,  the 
uterus  and  vagina,  and  in  different  divisions  of  the  male  genital  system. 
The  term  enteric  system  is  applied  to  this  type  of  nerve  structure,  specific- 
ally that  part  typical  of  the  alimentary  canal.  These  nets,  which  are  for 
the  most  part  microscopic,  are  also  freely  connected  with  other  parts  of 
the  system,  as  well  as  with  the  cerebro-spinal  axis. 

Gross  Anatomical  Relations. — No  special  interest  attaches  to  the  gross 
anatomical  relations  outside  of  the  fact  that  these  factors  are  guides  to  the 
complicated  neurone  relations  in  which  our  present  interest  in  the  autono- 
mic system  lies.  The  ganglia  of  the  sympathetic  chain  typically  corre- 
spond segment  by  segment  with  the  spinal  nerves  with  which  they  are 
connected.  The  type  arrangement  is  shown  in  figure  417.  Each  ganglion 
is  connected  with  its  corresponding  spinal  nerve  by  two  nerve  branches, 
the  white  and  gray  rami. 

The  secret  of  the  presence  of  the  two  kinds  of  rami  is  revealed  by  the 
work  of  Gaskell  on  the  type  and  character  of  the  constituent  neurones  of 
the  white  and  gray  rami  and  of  the  spinal  nerves,  and  of  Langley  and 
Anderson  on  the  neurone  connections  and  neurone  pathways.  It  has  been 


666  THE    NERVOUS    SYSTEM 

determined  that  the  cord  neurones  contributing  fibers  to  the  autonomic 
system  have  their  cell  bodies  of  origin  in  the  lateral  and  antero-lateral 
gray  columns  of  the  cord.  Their  axones  are  small  medullated  fibers  which 
compose  the  major  portion  of  the  white  rami.  They  are  a  third  or  a 
fourth  of  the  diameter  of  ordinary  medullated  fibers,  measuring  i.Sju 
to  2.7/x  instead  of  141*  to  19/4.  Such  fibers  are  peculiar  to  the  spinal  nerve 
roots  of  the  thoracic  group  and  of  the  upper  lumbar  nerves.  But  they  are 
also  found  in  the  second  and  third  sacral  nerves  and  constitute  there  the 
nervi  erigentes  which  pass  directly  to  the  hypogastric  plexus.  These  fibers 
end  in  arborizations  about  the  cell  bodies  of  the  sympathetic  ganglia. 

White  rami  are  lacking  in  the  entire  cranial  and  cervical  regions. 
They  are  absent  also  in  the  lumbo-sacral  cord  below  the  fourth  lumbar 
segment.  Since  the  superior,  middle,  and  inferior  cervical  ganglia 
are  the  only  cervical  representatives  of  the  chain  ganglia,  it  is  evident 
that  there  is  an  atypical  distribution  of  gray  rami  in  the  neck  and  head. 
The  gray  rami,  distributed  to  cervical  spinal  nerves,  arise  in  the  first 
thoracic  and  inferior  and  the  superior  cervical  ganglia. 

Sy.    G.  Pr.  G.  7h 


FIG.  418. — Langley's  autonomic  types  of  preganglionic,  black,  and  postganglionic, 
red,  neurones.  Sp.  C.  =  Spinal  cord.  Sy.  G  .  =  Sympathetic  ganglia.  Pr.  G.  =  Pe- 
ripheral ganglia,  solius,  hypogastric;  etc.  Tr.  =  Terminations  in  muscle,  gland,  etc. 


The  axones  of  the  cells  of  the  ganglia,  fibers  of  the  second  order,  con- 
stitute the  continuation  pathways  either  by  the  gray  rami  back  to  the 
somatic  nerves  and  on  to  a  distribution  in  the  blood  vessels,  and  the  glands 
and  skin  of  the  trunk  Or  they  often  run  by  direct  visceral  branches  to 
the  lungs,  heart,  alimentary  canal,  urogenital  system,  etc. 

The  autonomic  pathway,  from  the  histological  and  functional  stand- 
point, always  consists  of  a  two  neurone  chain.  The  last  link  in  this  chain 
is  called  by  Langley  the  postganglionic  neurone.  The  connecting  link 
between  the  cord  and  brain  stem  and  the  sympathetic  chain  ganglia  he 
designates  the  preganglionic  neurone.  Some  of  the  ganglionic  neurones 


THE    DOUBLE    SYSTEM  667 

do  not  form  synapses  immediately  they  enter  a  chain  ganglion  but  pass  on 
through  to  more  distant  ganglia,  Pr.  G.,  figure  418. 

The  determination  of  the  location  of  the  synapses  between  pre-  and 
postganglionic  neurones  was  accomplished  by^the  brilliant  work  of  Langley 
and  Anderson  with  the  nicotine  method.  They  used  the  discovery  that 
nicotine  readily  poisons  the  synapses  of  preganglionic  arborization  on 
postganglionic  neurones,  thus  effectively  blocking  physiological  conduction 
through  the  synapse. 

When  nicotine  is  injected  into  the  circulation  or  painted  directly  onto 
a  ganglion  it  at  first  stimulates,  then  completely  paralyzes  the  endings  of 
any  preganglionic  arborizations  about  the  cell  bodies  of  the  postganglionic 
neurones  of  the  group.  Nerve  fibers  as  such  do  not  lose  their  functions 
under  these  conditions.  This  observation  enables  one  by  the  use  of  the 
nicotine  method  to  determine  in  what  particular  ganglion  nerve  impulses 
to  a  peripheral  organ  are  blocked.  Langley  and  his  students  reexplored 
the  physiological  distribution  of  fibers  controlling  visceral  organs. 

They  found  for  example  that  the  sympathetic  nerves  for  the  head  and 
upper  portion  of  the  neck,  which  have  their  origin  in  the  second  to  the 
fifth  thoracic  spinal  nerve  roots,  have  their  pre-postganglionic  synapses 
in  the  superior  cervical  ganglion.  The  outflow  of  autonomic  nerves  to 
the  abdominal  viscera  which  occurs  in  the  cord  from  the  seventh  thoracic 
to  the  fourth  lumbar  have  their  preganglionic  terminations  in  the  ganglia 
of  the  celiac  and  the  mesenteric  plexuses.  The  pelvic  viscera  receive  their 
fibers  from  the  first  four  lumbar  spinal  nerves  and  the  synapses  are  in 
the  inferior  mesenteric  ganglia  and  certain  smaller  more  distal  ganglia. 
The  type  arrangement  is  diagrammatically  presented  by  Langley  as  shown 
in  figure  418. 

The  Double  System. — The  description  of  the  outflow  of  efferent  nerve 
fibers  from  the  central  nervous  axis  to  autonomic  mechanisms  has  thus 
far  been  based  on  the  type  region,  i.e.,  from  the  first  dorsal  to  the  fourth 
lumbar  inclusive.  However,  certain  other  regions  of  the  central  axis 
give  rise  to  nerve  fiber  groups  that  bear  both  anatomical  and  physiological 
similarities  to  the  dorsal  region.  These  regions  are  that  portion  of  the 
brain  stem  which  gives  rise  to  the  oculomotor  or  third  cranial  nerve,  espe- 
cially those  fibers  which  innervate  the  ciliary  muscle  and  the  iris ;  the  motor 
roots  of  the  seventh,  ninth  and  tenth  cranial  nerves  containing  secretory, 
vasodilator,  motor  and  inhibitory  nerve  fibers;  and  the  region  of  the 
third  and  fourth  segments  of  the  sacral  cord  which  gives  rise  to  the  fibers 
distributed  to  the  urogenital  system  and  muscles  of  the  lower  bowel.  In 
each  of  these  groups  of  anatomical  nerves  there  is  a  pre-  and  postgang- 
lionic neurone  in  the  physiological  path  to  the  terminal  tissue.  In  the 
third  cranial  nerve,  for  example,  the  ciliary  ganglion  is  the  location  of  the 
pre-postganglionic  synapse.  In  the  inhibitory  pathway  of  the  vagus  this 


668  THE    NERVOUS    SYSTEM 

synapse  takes  place  in  the  cardiac  ganglion.  In  short,  the  second  group 
of  ganglia  described  in  the  introductory  paragraph  on  the  autonomic 
system  belongs  to  nerve  mechanisms  of  this  special  group. 

The  efferent  nerve  groups  of  the  cranial,  thoracic  and  sacral  regions 
are  all  similar  in  one  functional  regard,  viz.,  there  is  no  voluntary  control 
of  the  functions  which  they  influence.  These  nerves  as  a  whole  innervate 
the  eyes,  salivary  glands,  blood  vessels,  heart,  bronchial  tubes,  stomach,  in- 
testine, colon,  liver,  pancreas,  kidney,  urinary  bladder  and  genital  organs, 
all  so-called  involuntary  organs  and  mechanisms.  All  of  these  organs 
have  a  double  innervation  as  shown  in  detail  in  figure  418.  This  simi- 
larity of  neurone  pathway  and  of  involuntary  functional  control  led  Lang- 
ley  to  reclassify  the  whole  nerve  outflow  under  the  name,  the  autonomic 
system. 

It  will  be  seen  that  each  visceral  organ  receives  groups  of  fibers  from 
two  quite  different  sections  of  the  brain  or  cord.  However,  physiologically 
these  two  sources  of  nerves  provide  the  organ  with  diametrically  opposed 
nerve  influences  controlling  or  regulating  its  function.  For  example, 
in  the  iris  of  the  eye  one  set  of  fibers  stimulates  constriction  of  the  pupil 
and  the  other  dilation.  One  set  of  fibers,  the  vagus,  inhibits  the  heart, 
the  other  set,  the  augmentors,  accelerates  it,  etc.  Autonomic  fibers  from 
the  thoracico-lumbar  cord  in  general  have  exactly  the  opposite  influence  on 
function  which  the  fibers  from  the  brain  stem  and  the  sacral  cord  exert. 
The  similarity  of  functional  differentiation  is  revealed  also  by  the  similarity 
of  action  of  drugs  on  the  cranial  and  sacral  nerves,  as  contrasted  with 
the  thoracic  and  lumbar  nerves.  This  group  of  autonomic  nerves  may 
therefore  be  subclassed  into  cranio-sacral  and  thoracico-lumbar  autonomies. 
Langley  originally  classified  the  oculomotor  and  medullary  outflow, 
and  the  sacral  outflow  as  parasympathetic,  in  contrast  with  the  thoracic, 
for  which  he  retains  the  name  sympathetic.  It  seems  better  to  use  the 
more  logical  classification  in  the  functional  grouping  as  outlined. 

To  the  more  diffuse  innervation  of  the  walls  of  hollow  organs,  such  as  the 
stomach  and  intestine,  Langley  gave  the  name  enteric  system,  see  page  669. 
The  best  known  portion  of  the  enteric  system  consists  of  the  myenteric 
plexus  (Auerbach)  and  the  submucous  plexus  (Meissner).  These  diffuse 
nets  of  ganglia  are  the  only  groups  of  nerve  cells  outside  of  the  central 
axis  in  the  mammalia  known  to  coordinate  reflexes.  The  system  is 
phylogenetically  an  older  nerve  differentiation.  The  enteric  ganglia  are 
also  pathways  for  extrinsic  autonomic  nerves  from  the  cerebrospinal  axis 
in  which  it  is  assumed  with  little  or  no  proof  that  the  enteric  ganglion  cells 
are  postganglionic  in  their  homologies.  Portions  of  the  urogenital  system, 
tubules  and  vesicles,  possess  a  similar  diffuse  nerve  mechanism.  The 
whole  schema  is  tabulated  below.  This  classification  is  now  rather 


THE    AUTONOMIC    NERVOUS    SYSTEM 


669 


broadly  accepted  in  physiology  and  forms  a  more  rational  system  around 
which  we  may  group  the  divers  physiological  functions,  see  figure  419. 

THE  AUTONOMIC  NERVOUS  SYSTEM. 


II. 


The  Thoracicolumbar  Autonomies. 

The  First  dorsal  to  the  Fourth  lumbar  inclusive. 
The  Craniosacral  Autonomies. 
The  midbrain  autonomic  nerves. 
The  3d  cranial  or  oculomotor. 
The  bulbar  autonomies. 
The  yth  or  facial. 
The  9th  or  glossopharyngeal. 


^x 


SubmaxittcLrV 

Parotid 
Head  blood 


,-  Somatic  vascular, 
/secretory  and  pilomotor 


Colon-rectum, 
Urinary  bladder 
Genital  organs 


FIG.  419. — Diagrammatic    representation   of   the    paths   of   the    autonomic   nervous 
distribution.     (Modified  from  Meyers  and  Gottlieb.) 

The  loth  or  vagus. 

The  nth  or  spinal  accessory. 
The  sacral  autonomies. 

The  2d  and  3d  sacral  or  the  nervus  erigens. 
III.  The  enteric  or  visceral  autonomies. 
Plexus  of  Auerbach. 
Plexus  of  Meissner. 


670  THE    NERVOUS    SYSTEM 

Functions  of  the  Autonomic  Nerves. — The  Thoracicolumbar  Autonom- 
ies.— As  we  have  already  seen,  the  thoracicolumbar  fibers  have  their  spinal 
origin  from  the  first  thoracic  to  the  fourth  lumbar  spinal  segments.  Yet 
these  fibers  are  distributed  to  all  portions  of  the  body.  They  accomplish 
the  following  functions :  vasomotors  to  the  blood  vessels  of  the  entire  body ; 
vasodilators  to  the  blood  vessels  of  the  body  exclusive  of  the  portions  of  the 
skin  of  the  head;  the  salivary  gland,  gastric  and  pancreatic  glands  and 
urogenital  system;  they  furnish  motor  nerves  to  the  heart  and  inhibitory 
nerves  to  the  bronchi,  stomach  and  intestine,  and  both  motor  and  inhibi- 
tory nerves  to  portions  of  the  urogenital  system;  secretory  nerves  for  the 
sweat  glands  throughout  the  body;  and  the  pilomotor  nerves  of  the  skin. 
The  details  of  functional  control  of  these  divisions  of  the  thoracic  auto- 
nomies have  been  discussed  in  presenting  the  physiology  of  the  organs 
concerned. 

The  Third  Cranial  Nerve. — Langley  has  divided  the  cranial  autonomies 
into  the  mid-brain  autonomies — the  third  cranial  nerves,  and  the  bulbar 
autonomies — the  seventh,  ninth,  tenth  and  eleventh  cranial  nerves. 

The  oculomotor  or  third  cranial  nerve  is  an  efferent  nerve  distributed 
to  the  extraocular  muscles  and  to  the  muscles  of  the  iris  and  ciliary  appara- 
tus. The  fibers  to  the  extraocular  muscles  are  of  the  usual  skeletal  motor 
type.  The  fibers  of  the  second  group  belong  to  the  autonomic  system. 
These  nerves  run  by  way  of  the  ciliary  ganglia  where  they  have  their  pre- 
postganglionic  synapses.  They  are  distributed  to  the  circular  muscles  of 
the  eyes  which  on  stimulation  cause  constriction  of  the  pupils,  and  to  the 
two  sets  of  muscles  in  the  ciliary  apparatus  which  on  stimulation  lead  to 
the  act  of  accommodation. 

The  Seventh  Cranial  Nerve. — The  intermediate  nerve  of  Wrisberg  con- 
tains the  autonomic  fibers  of  the  facial,  figure  413.  Fibers  of  this  group 
constitute  the  chorda  tympani  whose  function  was  discovered  by  Bernard 
when  he  demonstrated  the  presence  of  secretory  fibers  to  the  submaxillary 
gland.  He  also  proved  the  presence  of  vasodilator  nerves  through  this 
same  channel.  The  chorda  tympani  stands  today  as  the  typical  example 
of  a  vasodilator  and  secretory  nerve,  both  functions  of  the  involuntary  or 
autonomic  motor  type.  The  cell  bodies  of  the  postganglionic  neurones 
lie  in  the  ganglia  at  the  hilus  of  the  gland.  The  seventh  apparently  also 
distributes  preganglionic  fibers  to  the  spheno-palatine  ganglion  from 
whence  postganglionic  fibers  are  distributed  to  the  mucous  membrane  of 
the  nose  and  upper  respiratory  passages. 

The  Ninth  Cranial  Nerve. — The  tympanic  branch  of  the  ninth  cranial 
nerve  supplies  vasodilator  and  secretory  fibers  to  the  parotid  gland. 


FUNCTIONS    OF   THE   AUTONOMIC   NERVOUS    SYSTEM 


67I 


The  Tenth  and  Eleventh  Cranial  Nerves. — The  vagus  or  tenth  cranial 
nerve  is  perhaps  the  most  composite  of  all  the  autonomic  nerve  pathways. 
It  together  with  the  roots  distributed  to  it  from  the  accessory  supplies 
inhibitory  fibers  to  the  heart;  motor  fibers  to  the  bronchi,  esophagus, 
stomach  and  intestine;  and  secretory  fibers  to  the  gastric  glands,  pancreas 
and  liver.  The  pre-postganglionic  unions  are  found  in  small  terminal 
ganglia  such  as  the  local  cardiac  ganglia. 

The  Sacral  Autonomies. — These  fibers  arise  from  the  sacral  cord  at  the 
level  of  the  second  and  third  sacral  nerves.  The  fibers  do  not  join  the 


M 


S 


FIG.  4iqA. — Diagrammatic  representation  of  the  scheme  of  innervation  of  the  ali- 
mentary canal.  A,  mucosa;  B,  sensory  nerve  endings;  C,  gastric  or  intestinal  wall; 
D  and  E,  sensory  and  motor  cell  bodies  in  the  enteric  plexus;  M,  motor  neurons, 
vagus  fibers  for  the  stomach  and  upper  portion  of  the  intestine,  sympathetic  fiber 
lower  down;  S  and  S',  afferent  sensory  nerves;  Sy,  sympathetic  ganglion,  pre-post- 
ganglionic synapse,  inhibitory  path  for  the  mammalian  canal.  The  fiber  S'  is  introduced 
to  Dixon's  original  figure  on  the  basis  of  his  foot-note.  (Greene,  modified  from  Dixon.) 

main  sympathetic  chain  but  are  distributed  in  the  nervus  erigens  by  way 
of  the  ganglia  of  the  pelvic  plexus.  The  nerves  furnish  motor  fib,ers  to  the 
bladder,  the  descending  colon  and  rectum;  inhibitory  fibers  to  the  sphinc- 
ter muscles  of  the  bladder.  They  also  contain  vasodilators  for  the  rectum, 
anus  and  penis;  also  inhibitory  fibers  to  the  retractor  penis  muscle.  The 
urogenital  system  obviously  receives  its  double  innervation  in  part  from 
the  thoracic  autonomies  and  in  part  from  the  sacral  autonomies. 

The  Enteric  Nervous  System. — The  enteric  autonomies  consist  of  those 
ganglia  and  fibers  lying  between  the  muscle  walls  of  such  hollow  organs  as 
the  stomach,  intestine,  etc.  Among  all  the  numerous  peripheral  ganglia 
of  the  body  the  ganglia  in  the  stomach  and  intestinal  wall  are  the  only  ones 


672  THE    NERVOUS    SYSTEM 

for  which  we  have  proof  of  local  reflex  control.  The  regulation  of  the  py- 
loric  sphincter  through  the  acid  control  is  an  example  of  such  local  reflex. 
This  reaction  takes  place  in  the  isolated  organ.  In  certain  ways  the  enteric 
system  seems  to  be  most  primitive  in  structure,  comparable  to  the  nervous 
system  we  find  in  lower  organism,  such  as  anemones  and  medusae.  How- 
ever, the  ganglion  cells  of  this  system  have  been  proven  to  arise  from  the 
neural  groove  and  to  migrate  to  their  adult  location  during  embryonic 
development.  Out  of  the  investigations  as  to  the  function  of  these 
mechanisms  we  have  the  idea  of  control  represented  in  figure  41 9  A. 

According  to  Gaskell  the  functions  of  the  main  sympathetic  ganglia  are 
the  following:  i.  The  sympathetic  ganglia  are  aggregates  of  large  numbers 
of  multipolar  cells  around  which  the  medullated  fibers  of  the  white  rami  form 
synapses.  The  branches  of  the  cells  are  of  the  non-medullated  fiber  type. 
Thus  a  medullated  conduction  is  converted  in  the  ganglia  into  a  non-medul- 
lated path  beyond  the  ganglia.  2.  The  ganglion  cells  exercise  a  nutritive 
influence  over  the  tissues  to  which  their  fibers  are  distributed.  3.  The  num- 
ber of  preganglionic  fibers  entering  the  ganglia  is  not  so  great  as  that  leaving, 
since  the  cells  are  multipolar  (not  shown  in  the  schematic  figure  417).  This 
serves  to  multiply  the  influence  of  a  relatively  simple  efferent  preganglionic 
conduction  path  and  to  extend  it  over  a  larger  area  the  parts  of  which  are 
usually  acting  co-ordinatively. 

The  sympathetic  ganglia  are  not  nerve  centers  in  the  usual  sense.  It  is 
better  to  regard  them  merely  as  distributing  organs  in  which  impulses  of  cen- 
tral origin  and  comparatively  simple  type  are  distributed  over  relatively 
large  areas.  These  ganglia  do  not  possess  the  power  of  reflex  function 
except  in  the  enteric  system  as  noted  above.  A  type  of  pseudo-reflex 
has  been  described  depending  on  the  law  of  neurone  reaction.  But  it  is 
not  supposed  that  such  reflexes  occur  in  the  normal  animal. 

Afferent  Fibers  in  Sympathetic  Paths. — Afferent  or  sensory  fibers  of 
the  ordinary  spinal-root  ganglion  cells  are  present  in  the  sympathetic 
nerves  of  the  splanchnic  region.  These  fibers  are  the  distal  branches  of 
posterior  root  ganglion  cells  and  are  distributed  to  the  visceral  region  by 
way  of  the  white  rami  and  the  sympathetic  system.  These  fibers  have 
their  terminations  in  the  mucosa  of  the  alimentary  and  urogenital  system 
or  in  the  muscular  walls  of  these  organs.  Their  stimulation  arouses  the 
sensation  of  fullness  (Hertz)  that  characterizes  hollow  organs,  also  sensa- 
tions of  pain  and  temperature  as  shown  by  Carlson  for  the  stomach.  It 
is  the  stimulating  influence  of  operative  conditions  on  this  class  of  sensory 
fibers  that  plays  so  important  a  part  in  surgical  shock.  Modern  practices 
in  anesthesia  are  especially  devised  to  prevent  this  dangerous  state  of  de- 
pression. Such  afferent  paths  are  not  a  constituent  part  of  the  autonomic 
classification  of  Langley.  However,  Dogiel  has  described  a  type  of  true 
afferent  sympathetic  neurone.  Their  fibers  arise  from  cells  located  in  the 


THE    PHYSIOLOGY    OF    SLEEP  673 

sympathetic  ganglia,  and  pass  through  the  rami  communicantes,  to  end  by 
terminal  arborizations  in  the  spinal  root  ganglia,  chiefly  around  cells  of 
the  Dogiel  type.  The  number  and  significance  of  this  class  of  afferent 
neurones  remain  as  yet  uncertain.  Though  their  spinal  relations  suggest 
some  association  with  the  phenomenon  of  referred  pain. 


THE  PHYSIOLOGY  OF  SLEEP. 

All  parts  of  the  body  which  are  the  seat  of  active  change  require  periods 
of  rest.  The  alternation  of  work  and  rest  is  a  necessary  condition 
of  their  maintenance  and  of  the  healthy  performance  of  their  functions. 
These  alternating  periods,  however,  differ  much  in  duration  in  different 
organs  or  regions  of  the  body.  But,  for  any  individual  instance  they  pre- 
serve a  general  and  rather  close  uniformity.  Thus,  the  periods  of  rest  and 
work  mentioned  in  the  case  of  the  heart  occupy,  each  of  them,  about  half  a 
second.  In  the  case  of  the  ordinary  respiratory  muscles  the  periods  are 
about  four  or  five  times  as  long.  In  many  cases  (as  of  the  voluntary 
muscles  during  violent  exercise) ,  while  the  periods  during  active  exertion 
alternate  very  frequently,  yet  the  expenditure  goes  far  ahead  of  the  repair, 
and  to  compensate  for  this  an  after-repose  of  some  hours  becomes  neces- 
sary, the  rhythm  being  less  perfect  as  to  time  than  in  the  case  of  the 
muscles  concerned  in  circulation  and  respiration. 

Obviously,  short  periods  of  activity  and  repose,  or  in  other  words,  of  con- 
sciousness and  unconsciousness  would  be  impossible  in  the  case  of  the  brain. 
The  repose  must  occur  at  long  intervals  and  must  be  proportionately  long. 
Hence  the  necessity  for  that  condition  which  we  call  Sleep;  a  condition 
which,  at  first  sight  seeming  exceptional,  is  only  an  unusually  perfect  exam- 
ple of  what  occurs  at  varying  intervals  in  every  actively  working  portion 
of  our  bodies. 

By  exposing  the  surface  of  the  brain  of  a  living  animal  at  a  circum- 
scribed spot,  and  protecting  the  exposed  part  by  a  watch-glass,  Durham 
was  able  to  prove  that  the  brain  becomes  visibly  paler,  anemic,  during 
sleep.  And  the  anemia  of  the  optic  disc  during  sleep,  observed  by  Hugh- 
lings  Jackson,  may  by  analogy  be  taken  as  a  strong  confirmation  of  the 
same  fact. 

The  Circulation  During  Sleep. — Blood  is  supplied  to  the  brain  in  four 
distinct  but  anastomosing  arteries.  This  efficient  anatomical  arrange- 
ment is  obviously  all  the  more  important  when  it  is  remembered  that  the 
circulation  in  the  brain  has  only  an  inefficient  local  device  for  regulating 
the  blood-flow,  and  that  the  circulation  of  the  brain  is  constantly  influ- 
enced by  the  variations  in  general  blood  pressure,  see  page  242. 

Howell  and  others  have  studied  the  circulation  by  the  plethysmographic 
method  during  sleep.  The  results  show  that  with  the  loss  of  consciousness, 

43 


674  THE    NERVOUS    SYSTEM 

and  immediately  following,  there  is  a  sharp  dilatation  of  the  blood  vessels  of 
the  arm,  probably  chiefly  of  the  skin,  as  shown  by  the  increase  in  volume. 
The  vessels  remain  dilated  until  the  individual  begins  to  awaken,  then  there 
is  a  rapid  constriction  with  decrease  of  volume  of  the  organ. 

The  dilatation  of  the  general  blood  vessels  draws  off  the  supply  of  blood 
from  the  brain,  and  the  resulting  partial  anemia  contributes  to  loss  of  con- 
sciousness. The  blood  supply  is  ample  for  the  growth,  repair  and  rest  of  the 
nervous  system.  How  efficient  this  rest  period  is  for  the  rejuvenation  of  the 
nervous  tissue  is  indicated  even  by  the  relatively  gross  means,  figure  357, 
shown  in  the  histological  preparations  of  nerve  cells. 

Somnambulism  and  Dreams. — What  we  term  sleep  occurs  often  in  very 
different  degrees  in  different  parts  of  the  nervous  system;  and  in  reference  to 
some  parts  the  expression  cannot  be  used  in  the  ordinary  sense.  For  exam- 
ple, during  the  most  profound  cerebral  sleep  the  medulla  is  discharging 
rhythmic  nerve  impulses  to  maintain  respiratory  movements. 

The  phenomena  of  dreams  and  somnambulism  are  examples  of  differing 
degrees  of  sleep  in  different  parts  of  the  cerebro-spinal  nervous  system.  In 
the  former  case  the  cerebrum  is  still  partially  active;  but  the  activity  is  no 
longer  corrected  by  the  reception,  on  the  part  of  the  sleeping  sensorium,  of 
impressions  of  objects  belonging  to  the  outer  world.  Neither  can  the  cere- 
brum in  this  half-awake  condition  control  the  centers  of  reflex  action  of 
the  voluntary  muscles  so  as  to  cause  the  latter  to  contract  in  close  co- 
ordination with  the  changing  cerebral  reactions  as  during  waking  hours 
— a  fact  within  the  painful  experience  of  all  who  have  suffered  from 
nightmare. 

In  somnambulism  the  higher  centers  are  capable  of  co-ordinating  that 
train  of  reflex  nervous  action  which  is  necessary  for  progression,  while  the 
nerve  center  of  the  muscular  equilibrium  sense  (in  the  cerebellum)  is,  pre- 
sumably, fully  awake;  but  the  sensorium  is  still  asleep,  and  impressions  made 
on  it  are  not  sufficiently  felt  to  rouse  the  cerebrum  to  a  comparison  of  the 
difference  between  mere  ideas  or  memories  and  sensations  derived  from 
external  objects. 


LABORATORY    EXPERIMENTS    ON    THE    NERVOUS    SYSTEM         675 

LABORATORY  EXPERIMENTS  ON  THE  NERVOUS  SYSTEM. 

1.  Irritability  of  the  Neurone. — Repeat  the  tests  outlined  in  Experi- 
ment 3  under  Laboratory  Experiments  in  Muscle.     Apply  these  tests  both 
to  nerve  trunk  and  to  any  ganglion,  for  example,  the  ganglion  on  the 
posterior  root  of  the  tenth  spinal  nerve  of  the  frog,  or  the  superior  cervical 
or  the  celiac  ganglion  of  the  cat. 

2.  Conductivity  of  the  Neurone.  — a.  Conductivity  can  be  demonstrated 
quantitatively  by  the  method  of  Experiment  19,  under  Muscle  Nerve 
Physiology,  using  the  muscle  as  an  index  of  nerve  activity. 

b.  A  better  method  is  to  measure  the  latency  of  the  action  current  of  an 
isolated  sciatic  nerve.     Connect  one  end  of  the  nerve  by  means  of  non- 
polarizable    electrodes   with    a    delicate   galvanometer.     Stimulate    the 
nerve  at  the  far  end,  then  at  the  near  end. 

c.  That  conduction  is  in  either  direction  can  also  be  demonstrated  on 
branched  motor  nerves.     Dissect  out  the  sartorius  muscle  of  the  frog  with 
its  motor  nerve,  which  has  two  branches.     Split  the  muscle  between  the 
branches.     Pick  up  one  end  and  stimulate  its  nerve  branch.     Both  muscles 
will  contract.     The  nerve  impulse  developed  at  the  point  of  stimulation 
also  travels  away  from  its  muscle  to  the  point  of  branching  of  the  axone, 
then  down  the  second  nerve  branch  to  stimulate  the  second  division  of  the 
muscle.     Branched  axones  are  involved,  not  separate  neurones.     This  is 
the  type  of  reaction  in  a  pseudo  reflex. 

3.  Differentiation  Polarity  in  Neurones. — Neurones  have  a  polarity 
because  of  two  facts,  first,  their  anatomical  relations,  and  second,  their 
physiological  contacts.     Afferent  or  sensory  nerves  cannot  be  stimulated 
normally  except  at  the  sense  organs.     The  nerve  impulses  developed  must 
pass  in  the  centripetal  direction,  affecting  such  other  neurones  as  are  in 
physiological  contact,  until  a  motor  organ  is  reached. 

Differentiations  can  be  demonstrated  by  certain  drugs,  for  example, 
paint  a  nerve  trunk  with  nicotine.  No  effect  follows.  Paint  a  ganglion, 
the  nicotine  stimulates,  then  poisons,  the  cell  bodies  of  the  ganglion.  Or, 
paint  the  accelerator  nerve  trunks  to  the  heart  with  o.i  per  cent,  adrenalin, 
no  obvious  change  occurs.  Paint  the  heart  or  perfuse  the  heart  with 
adrenalin  to  bring  the  hormone  in  contact  with  the  cardiac  motor  nerve 
endings.  The  endings  are  specifically  stimulated  and  the  heart  beats 
faster  and  stronger. 

4.  The  Afferent  and  Efferent  Functions  of  the  Spinal  Nerve  Roots.— 
Skilfully  dissect  and  expose  the  two  roots  of  the  eighth,  ninth,  and  tenth 
spinal  nerves  of  the  right  side  of  a  large  frog.     Reserve  the  left  side  for  a 
second  try.     Insert  silk  threads  under  each  root  using  care  not  to  stretch 
or  injure  the  delicate  nerves.     The  posterior  root  is  the  smaller  and  its 
orange-colored  ganglion  aids  in  identifying  it. 


676  THE    NERVOUS    SYSTEM 

a.  Stimulate  the  posterior  root  with  weak  induction  currents.     Vary 
the  strength  till  muscular  contractions  occur  in  the  leg  on  the  same  side. 

b.  Section   the   root   in   the   middle.     Stimulate   the  peripheral  end. 
Stimulate  the  central  end.     Do  not  allow  escape  of  the  stimulating  current 
to  other  tissues  nor  to  the  anterior  root. 

c.  Stimulate  the  anterior  root  and  note  the  contractions  that  result. 

d.  Section  the  anterior  root  in  its  middle  and  stimulate  the  peripheral 
end,  using  as  weak  a  stimulus  as  you  can  for  effective  contractions.     Now 
stimulate  the  central  end. 

e.  Demonstrate   the   crossed   reflex   if   possible   by   stimulating    the 
central  end  of  a  posterior  root  strongly.     Stimulate  the  central  end  of 
the  brachial  nerve  and  look  for  crossed  reflexes. 

/.  Expose  the  third  spinal  nerve,  cut  it  and  stimulate  the  central 
end  in  an  attempt  to  show  both  crossed  and  descending  reflexes  that  shall 
include  movements  of  the  hind  legs. 

5.  The  Reflex  Act,  Spinal  Frog. — Prepare  a  spinal  frog  by  transecting 
the  cord  where  it  joins  the  medulla,  the  posterior  border  of  the  tympanum, 
and  pith  the  brain.     This  spinal  animal  will  remain  quiet  in  whatever 
position. 

a.  Place  in  the  prone  position  with  the  hind  legs  stretched  full  length, 
pinch  the  tip  of  the  toe,  after  a  short  latent  period  the  leg  of  that  side  will 
be  drawn  up  into  the  usual  position  by  its  side.     If  the  stimulation  is 
continued  and  strong  enough  contractions  of  the  opposite  muscles  will 
occur. 

b.  Suspend  the  frog  by  its  jaws  and  stimulate  different  points  of  the 
skin  with  4  per  cent,  sulphuric  acid  applied  by  wet  bits  of  blotting  paper. 
Complicated  though  orderly  contractions  occur  in  the  muscles  of  now  one, 
now  the  other,  or  both  legs  according  to  which  spot  is  stimulated  by  the 
acid. 

c.  Stimuli  applied  in  the  median  line  of  the  body  lead  to  symmetrical 
contractions.     These  contractions  fail  if  the  spinal  cord  is  destroyed. 
Hence  they  are  reflex  acts,  simple,  or  coordinated;  unilateral  or  crossed, 
etc.     Convulsive  reactions  occur  when  the  stimulation  is  too   violent. 

6.  Decerebrate  Frog. — Prepare  a  series  of  frogs  by  destroying  portions 
of  the  brain  as  follows: 

a.  The  upper  portion  of  the  cerebrum,  not  injuring  the  olfactory  or 
optic  tracts. 

b.  All  the  cerebrum  without  injury  to  the  optic  lobes  and  tracts. 

c.  Destroy  the  cerebrum  and  optic  lobes,  without  injury  to  the  base 
of  the  brain  or  optic  tracts. 

d.  Destroy  all  parts  of  the  brain  in  front  of  the  medulla. 

e.  Section  at  the  point  of  union  of  the  spinal  cord  and  medulla  and 
destroy  the  brain  completely. 


LABORATORY   EXPERIMENTS    ON    THE    NERVOUS    SYSTEM         677 

Compare  this  series  of  frogs  with  the  behavior  of  the  normal  frog  as 
regards  the  following  points:  a.  position;  b.  spontaneous  movements; 
c.  movements  in  response  to  stimuli  of  different  types  and  intensities, 
i.e.,  electrical,  chemical,  disturbed  position,  etc.  These  notes  should  be 
carefully  tabulated  and  reported. 

7.  The  Decerebrate  Pigeon. — Observe  the  behavior  of  a  young  pigeon, 
then  remove  the  entire  cerebral  cortex.     The  operation  must  be  done 
aseptically  under  ether  and  Moo  grain  of  atropine.     Scoop  the  cortex 
out  with  care  not  to  injure  the  brain  ganglia,  optic  lobes  and   tracts. 
Control  bleeding  by  hot  tampons.     Close  the  wound  aseptically. 

Keep  the  animal  for  several  weeks,  feeding  and  watering  by  placing  food 
in  the  back  of  the  mouth.  Healing  will  quickly  occur  and  the  animal 
will  show  certain  normal  behaviors  from  the  moment  it  recovers  from 
ether.  Examine: 

a.  Its  spontaneous  activity. 

b.  Its  ability  to  take  feed,  swallow,  etc. 

c.  Its  responses  to  light,  mechanical  and  other  stimuli. 

d.  Its  ability  to  balance  on  a  perch  and  in  the  air. 

e.  Any  daily  improvement  in  its  reactivity. 

8.  Stimulation  of  the  Cerebral  Cortex  in  the  Dog  or  Cat. — Sensory  and 
motor  localizations  of  the  cortex  of  the  mammal  have  been  described  for 
certain  areas.     Under  chloral  and  ether  anesthesia  expose  the  anterior 
surface   of   the   cerebral   hemisphere   of   the   dog.     Identify   the   crucial 
fissure.     Stimulate  points  anterior,  lateral  and  posterior  to  this  fissure, 
recording  and  labeling  the  exact  points  on  a  map  prepared  for  the  purpose. 
Keep  records  of  all  muscular  reactions.     Areas  can  be  identified  which  on 
stimulation  produce  contractions  of  muscles  of  the  fore  leg,  of  the  hind  leg, 
or  of  various  parts  of  the  trunk  and  tail,  etc.,  verify  such  areas.     Stimulate 
for  short  periods  only.     Avoid  fatigue  and  protect  the  area  from  cooling. 
Especial  care  must  be  given  to  the  degree  of  anesthesia. 

9.  Spinal  Transection  and  Motor  Control. — Under  chloral  and  ether 
and  with  aseptic  technic  transect  the  spinal  cord  of  a  female  dog  at  the 
level  of  the  thirteenth  dorsal.     Aseptically  close  the  wound  and  take 
hospital  care  of  the  animal  while  it  is  under  observation. 

a.  All  voluntary  movements  of  the  hind  legs  and  pelvic  musculature 
cease.     It  may  be  necessary  to  support  the  hind  quarters. 

b.  Certain  reflexes  still  take  place  through  the  isolated  lumbar  seg- 
ments of  the  cord,  for  example,  micturition  and  defecation. 

c.  Vasomotor  control  is  at  first  lost,  but  is  slowly  regained  in  a  few 
days. 

10.  Spinal  Hemisection. — Expose  the  spinal  cord  under  chloral  and 
ether.     Carefully  hemisect  the  right  half  of  the  cord  in  the»  twelfth  seg- 
ment and  close  as  in  the  preceding  test.     Use  surgical  and  aseptic  technic. 

a.  Carefully  map  the  area  and  degree  of  analgesia. 


678  THE    NERVOUS    SYSTEM 

b.  Analyze  and  record  the  extent  of  motor  paralysis. 

c.  Analyze  all  other  disturbances. 

1 1 .  Spinal  Analgesia. — Temporary  effects  similar  to  complete  section 
of  the  cord  are  produced  on  injecting  cocaine  into  the  spinal  canal  at  the 
same  level.     Inject  i  cc.  of  2  per  cent,  cocaine  hydrochlorate  into  the 
spinal  canal  of  the  twelfth  dorsal  segment  of  a  dog.     Complete  analgesia 
and  loss  of  motor  control  occur  in  the  hind  legs  and  pelvic  parts.     This 
condition  passes  off  in  20  to  40  minutes  but  is  adequate  for  major  surgical 
operations. 

12.  Reflex  Reaction  Time  of  Man. — Arrange  an  electric  buzzer  signal 
in  circuit  with  a  lead  off  from  a  sixty  cycle  alternating  current  lighting 
system.     The  circuit  is  controlled  by  a  spring  key  with  pin  pricking 
attachment.     Use  a  rapid  moving  Harvard  drum  for  recording.     No 
time  signal  is  needed,  since  the  buzzer  not  only  records  the  opening  and 
closing  of  the  key  but  also  the  duration  of  the  latent  period  in  K20 
seconds.     Have  the  reagent  close  the  key  quickly  with  his  index  ringer. 
The  resulting  pinprick  will  be  sufficient  stimulus  to  produce  reflex  removal 
of  the  finger,  particularly  if  he  is  left  in  ignorance  of  the  pin.     An  electric 
shock  can  readily  be  arranged  to  stimulate  the  sensory  surface  only. 

13.  Voluntary  Reaction  Time  Conditioned  by  Simple  Choice. — Use  the 
signal  and  recording  apparatus  arranged  as  outlined  above.     For  the  pin 
substitute  a  simple  spring  key  with  flag  signal  device  arranged  to  be  used 
or  not  at  the  discretion  of  the  experimentor.     The  reagent  is  instructed 
to  close  the  key  and  open  it  instantly  provided  he  sees  the  flag  signal. 
Read  off  the  reaction  time  from  the  buzzer  vibrations.     Any  other  sensory 
channel  may  be  chosen  instead  of  sight. 

In  extended  experimental  work  more  and  more  complicated  nerve 
reactions  should  be  measured. 


CHAPTER  XV. 
THE  SENSES. 

THROUGH  the  medium  of  the  nervous  system  man  obtains  a  knowledge 
of  the  existence  both  of  the  various  parts  of  his  body  and  of  the  external 
world.  This  knowledge  is  based  upon  sensations  resulting  from  the  stimula- 
tion of  certain  centers  in  the  brain  by  nerve  impulses  conveyed  to  them  by 
afferent  nerves.  Under  normal  circumstances  the  following  structures, 
are  necessary  for  the  physical  development  and  mental  perception  of  a 
sensation:  a,  A  peripheral  organ  for  the  reception  of  the  impression;  b,  a 
nerve  for  conducting  it;  c,  a  nerve  center  for  feeling  or  perceiving  it. 

The  senses  may  be  conveniently  classified  according  to  the  sensation  which 
is  experienced.  Each  sense  organ  when  stimulated  is  supposed  to  lead 
to  a  sensation  of  distinctive  character.  Yet  many  of  the  sensations  are 
vaguely  defined,  such  as  thirst,  fatigue,  etc.  Other  sensations  have  a  very 
definite  and  readily  identified  quality  such  as  sight,  taste,  etc.  Sensations, 
whether  definite  or  vague,  are  referred  by  us  to  some  source  or  origin  either 
in  the  body,  i,  the  internal  or  so-called  common  sensations,  or  outside  the 
body,  2,  the  external  or  special  senses.  No  sharp  line  can  be  drawn  in 
this  classification. 

Internal  Senses. — Under  this  head  fall  all  those  senses  which  produce 
sensations  that  are  referred  to  an  origin  within  the  body,  such  as  fatigue, 
discomfort,  faintness,  satiety,  nausea,  together  with  hunger,  thirst,  the  muscle 
sense,  and  pain.  In  hunger  and  thirst  there  is  a  general  bodily  discomfort, 
but  in  many  persons  also  a  distinct  sensation  referred  to  the  stomach  or  to 
the  fauces.  In  this  class  must  also  be  placed  the  various  stimulations  of 
the  mucous  membrane  of  the  bronchi,  which  give  rise  to  coughing,  and  also  the 
sensations  derived  from  various  viscera.  It  is  by  means  of  the  muscle  sense 
that  we  become  aware  of  the  condition  of  the  muscles,  and  thus  obtain  the 
information  necessary  for  their  adjustment  to  various  purposes — standing, 
walking,  grasping,  etc.  This  muscular  sensibility  is  shown  in  our  power  to 
estimate  the  differences  between  weights  by  the  different  muscular  efforts 
necessary  to  raise  them.  It  must  be  carefully  distinguished  from  the  sense 
of  contact  or  pressure,  of  which  the  skin  is  the  organ.  When  standing  erect, 
we  can  feel  the  ground  contact,  and  there  is  a  sense  of  pressure,  due  to  our 
feet  being  pressed  against  the  ground  by  the  weight  of  the  body.  Both 
these  are  derived  from  the  skin  of  the  sole  of  the  foot.  If  now  we  raise  the 
body  on  the  toes,  we  are  conscious,  through  the  muscular  sense,  of  a  muscular 

679 


680  THE    SE.NSES 

effort  made  by  the  muscles  of  the  calf.  But  the  muscle  sense  will  be  dis- 
cussed further,  page  687. 

It  is  manifestly  impossible  to  draw  a  very  clear  line  of  demarcation 
between  some  of  these  senses. 

Special  Senses. — The  special  senses  include  Touch,  Temperature 
(Heat  and  Cold),  Taste,  Smell,  Hearing,  Sight. 

The  most  important  distinction  between  common  and  special  sensations 
is  that  by  the  former  we  are  made  aware  of  certain  conditions  of  various 
parts  of  our  bodies,  while  from  the  latter  is  gained  a  knowledge  of  the  ex- 
ternal world.  This  difference  will  be  clear  if  we  compare  the  sensations  of 
pain  and  touch,  the  former. of  which  is  a  common,  the  latter  a  special,  sensa- 
tion. "If  we  place  the  edge  of  a  sharp  knife  on  the  skin,  we  feel  the  edge 
by  means  of  our  sense  of  touch;  we  perceive  a  sensation,  and  refer  it  to  the 
object  which  has  caused  it.  But  as  soon  as  we  cut  the  skin  with  the  knife, 
we  feel  pain,  a  feeling  which  we  no  longer  refer  to  the  cutting  knife,  but  which 
we  feel  within  ourselves,  and  which  communicates  to  us  the  fact  of  a  change 
of  condition  in  our  own  body.  By  the  sensation  of  pain  we  are  neither  able 
to  recognize  the  object  which  caused  it  nor  its  nature. " 

It  is  important  in  studying  the  phenomena  of  sensation  clearly  to  under- 
stand that  the  sensorium,  or  seat  of  sensation,  is  in  the  brain,  and  not  in  the 
particular  organ  through  which  the  sensory  impression  is  received.  In  com- 
mon parlance  we  are  said  to  see  with  the  eye,  hear  with  the  ear,  etc.,  but  in 
reality  these  organs  are  only  adapted  to  receive  stimuli  which  produce  changes 
that  are,  through  their  respective  nerves,  conducted  to  the  sensorium,  to 
give  rise  to  sensation  of  sight,  hearing,  etc. 

Hence,  if  the  optic  nerve  is  severed,  vision  is  no  longer  possible.  Although 
the  image  falls  on  the  retina  as  before,  the  sensory  impulse  can  no 
longer  be  conveyed  to  the  sensorium.  When  any  given  sensation  is  felt,  all 
that  we  can  with  certainty  affirm  is  that  some  part  of  the  brain  is  excited. 
The  exciting  cause  may  be  some  object  of  the  external  world,  producing  an 
objective  sensation;  or  the  condition  of  the  sensorium  may  be  due  to  some 
excitement  within  the  brain  itself,  in  which  case  the  sensation  is  termed  sub- 
jective. The  mind  habitually  refers  sensations  to  external  causes;  and  hence, 
whenever  they  are  subjective  we  can  hardly  divest  ourselves  of  the  idea  of  an 
external  cause,  and  an  illusion  is  the  result. 

Sensory  Illusions. — Numberless  examples  of  such  illusions  might 
be  quoted.  As  familiar  cases  may  be  mentioned  humming  and  buzzing  in 
the  ears  caused  by  some  irritation  of  the  auditory  nerve  center.  These 
stimuli  may  even  be  interpreted  as  musical  sounds,  or  voices,  sometimes 
termed  auditory  spectra.  So-called  optical  illusions  in  which  objects  are 
described  as  seen,  although  not  present,  may  be  caused  by  changes  going 
on  in  some  part  of  the  visual  apparatus  beyond  the  eye.  Such  illusions  are 
most  strikingly  exemplified  in  cases  of  delirium  tremens  or  other  forms  of 


THE    SENSE    OF    TOUCH  68 1 

delirium,  and  may  take  the  form  of  animals  such  as  cats,  rats,  or  creeping 
loathsome  forms,  etc. 

One  uniform  internal  cause,  which  may  act  on  all  the  nerves  of  the  senses 
in  the  same  manner,  is  capillary  congestion.  This  one  cause  excites  in  the 
retina,  while  the  eyes  are  closed,  the  sensations  of  light  and  luminous  flashes; 
in  the  auditory  nerve,  the  sensation  of  humming  and  ringing  sounds;  in  the 
olfactory  nerve,  the  sense  of  odors;  and  the  nerves  of  feeling,  the  sensation 
of  pain.  In  the  same  way  a  chemical  substance  introduced  into  the  blood 
may  excite  in  the  nerves  of  each  sense  peculiar  symptoms:  In  the  optic  nerves, 
the  appearance  of  luminous  sparks  before  the  eyes;  in  the  auditory  nerves, 
tinnitus  aurium;  and  in  the  common  sensory  nerves,  the  sensations  of  creeping 
over  the  surface.  So,  also,  among  external  causes,  the  stimulus  of  electricity, 
or  the  mechanical  influence  of  a  blow,  concussion,  or  pressure,  excites  in  the 
eye  the  sensation  of  light  and  colors;  in  the  ear,  a  sensation  of  a  sound 
or  of  ringing;  and  in  the  tongue,  a  saline  or  acid  taste. 

Sense  Perceptions. — The  habit  of  constantly  referring  our  sensa- 
tions to  external  causes  leads  us  to  interpret  the  various  modifications  which 
external  objects  produce  in  our  sensations,  as  properties  of  the  external  bodies 
themselves.  Thus  we  speak  of  certain  substances  as  possessing  a  disa- 
greeable taste  and  smell;  whereas,  the  fact  is  their  taste  and  smell  are  only 
disagreeable  to  us.  It  is  evident,  however,  that  on  this  habit  of  referring  our 
sensations  to  causes  outside  ourselves,  perception,  depends  the  reality  of 
the  external  world  to  us;  and  more  especially  is  this  the  case  with  the  senses 
of  touch  and  sight.  By  the  co-operation  of  these  two  senses,  aided  by  the 
others,  we  are  enabled  gradually  to  attain  a  knowledge  of  external  objects 
which  daily  experience  confirms,  until  we  come  to  place  unbounded  confi- 
dence in  what  is  termed  the  evidence  of  the  senses. 

We  must  draw  a  distinction  between  mere  sensations,  and  the  judgments 
based,  often  unconsciously,  upon  them.  Thus,  in  looking  at  a  near  object, 
we  unconsciously  estimate  its  distance  and  say  it  seems  to  be  ten  or  twelve 
feet  off.  But  the  estimate  of  its  distance  is  in  reality  a  judgment  based  on 
many  things  besides  the  appearance  of  the  object  itself;  among  which  may 
be  mentioned  the  number  of  intervening  objects  and  their  relative  size, 
the  number  of  steps  which  from  past  experience  we  know  we  must  take 
before  we  can  touch  it,  etc. 

I.  THE  SENSES  OF  TOUCH,  TEMPERATURE,  PAIN,  AND 
THE  MUSCLE  SENSE. 

The  Sense  of  Touch. — The  sense  of  touch,  like  all  the  special  senses, 
possesses  a  special  end-organ  for  the  initiation  of  a  nerve  impulse  in  this 
instance  through  the  stimulus  of  contact  with  external  objects.  The  sense 
organ  of  touch  is  not  confined  to  particular  parts  of  the  body  of  small  extent, 


682  THE    SENSES 

like  the  organ  of  sight,  for  example,  but  is  found  in  all  parts  of  the  skin  and  its 
inversions-,  the  stomodeum  and  proctodeum.  The  nerves  of  touch  sensa- 
tion are  contained  in  the  same  trunks  with  other  sensory  nerves.  They  are 
found  in  the  posterior  or  sensory  roots  of  the  spinal  nerves  and  in  the  sensory 
divisions  of  the  cranial  nerves,  especially  the  fifth,  seventh,  ninth,  and  tenth. 
All  parts  of  the  epidermis  supplied  with  sensory  nerves  are  thus,  in  some 
degree,  organs  of  touch,  yet  the  sense  is  exercised  in  greatest  perfection  in 
certain  parts,  the  sensibility  of  which  is  extremely  delicate,  e.g.,  the  skin  of 
the  hands,  the  tongue,  and  the  lips,  which  are  provided  with  abundant  touch 
papillae.  A  peculiar  and  very  acute  sense  of  touch  is  exercised  through  the 
medium  of  the  nails  and  teeth,  and,  to  a  less  extent,  the  hair  may  be  consid- 
ered an  organ  of  touch,  as  in  the  case  of  the  eyelashes.  It  has  been  computed 
that  the  human  body  possesses  over  500,000  touch  spots. 


FIG.  420. — Touch  Corpuscle. 

The  sense  of  touch  renders  us  conscious  of  the  presence  of  a  contact 
stimulus,  from  the  slightest  to  the  most  intense  degree  of  its  action.  The 
modifications  of  this  sense  often  depend  on  the  extent  of  the  parts  affected. 
The  sensation  of  pricking,  for  example,  is  produced  when  the  sensitive  fibers 
are  intensely  affected  in  a  small  extent;  the  sensation  of  pressure  indicates 
a  slighter  affection  of  the  parts  over  a  greater  extent  and  depth.  It  is  by  the 
depth  to  which  the  parts  are  affected  that  the  feeling  of  pressure  is  distin- 
guished from  that  of  mere  contact. 

In  almost  all  parts  of  the  body  which  have  delicate  tactile  sensibility  the 
epidermis,  immediately  over  the  dermal  papillae,  is  moderately  thin.  When 
its  thickness  is  much  increased,  as  over  the  heel,  the  sense  of  touch  is  very 
much  dulled.  On  the  other  hand,  when  it  is  altogether  removed,  and  the 


ACUTENESS    OF    THE    SENSE  683 

cutis  laid  bare,  the  sensation  of  contact  is  replaced  by  one  of  pain.  Further, 
in  all  highly  sensitive  parts,  the  papillae  are  numerous  and  highly  vascular, 
and  the  sensory  nerves  are  connected  with  special  end-organs  which  have 
been  described  on  page  74  et  seq. 

The  special  endings  of  the  nerves  which  have  to  do  with  touch  may, 
however,  be  again  mentioned  here.  They  are  of  two  kinds,  viz.:  i.  Touch 
corpuscles,  which  are  found  chiefly  in  the  hands  and  feet,  particularly  on 
the  palmar  surface  of  the  hands  and  fingers,  but  also  on  the  under  sXirface 
of  the  forearm,  on  the  nipple,  eyelids,  lips,  and  the  genital  organs.  Touch 
corpuscles  are  situated  in  the  cutis  vera.  2.  End  bulbs  are  found  in  the 
conjunctiva  and  other  mucous  membranes,  the  lips,  genital  organs,  tongue, 
rectum,  and  elsewhere,  but  not  in  the  skin  proper.  As  regards  the 
Pacinian  corpuscles  and  similar  end-organs^  which  are  so  widely  distrib- 
uted, and  which  may  be  in  some  way  connected  with  the  development  of 
the  sensation  of  touch,  when  they  are  found  in  the  skin  they  are  situated 
very  deeply  in  the  cutis  vera  or  in  the  subcutaneous  tissue.  They  are 
extremely  numerous  on  the  nerves  of  the  palmar  surface  of  the  fingers. 
In  addition  to  these  special  nerve  endings  in  sense  organs,  nerve  fibers 
terminate  everywhere  in  the  skin  between  the  cells  of  the  Malpighian 
stratum  of  the  epidermis. 

The  acuteness  of  the  sense  of  touch  depends  in  no  small  degree  on  the 
cutaneous  circulation  and  is  of  course  greatly  influenced  by  external  tem- 
perature. This  explains  the  numbness  familiar  to  everyone  that  is  pro- 
duced by  the  application  of  cold  to  the  skin. 

Acuteness  of  the  Sense. — The  perfection  of  the  sense  of  touch  on 
different  parts  of  the  surface  is  proportional  to  the  minimal  pressure  re- 
quired to  stimulate  the  point,  i.e.,  the  threshold  stimulus.  Or  it  can  be 
measured  by  the  power  which  such  parts  possess  of  distinguishing  and 
isolating  the  sensations  produced  by  two  stimulating  points  placed  close 
together.  This  latter  is  in  a  degree  a  measure  of  the  power  of  localiza- 
tion. This  power  depends,  at  least  in  part,  on  the  number  of  primitive 
nerve  fibers;  for  the  fewer  the  primitive  fibers  which  an  organ  receives,  the 
more  likely  is  it  that  several  impressions  on  different  contiguous  points  will 
act  on  only  one  nerve  fiber,  and  hence  be  confounded,  and  perhaps  produce 
but  one  sensation.  Experiments  have  been  made  to  determine  the 
tactile  properties  of  different  parts  of  the  skin,  as  measured  by  this  power 
of  distinguishing  distances  between  points  of  simultaneous  contact. 
These  consist  in  touching  the  skin  with  the  points  of  a  pair  of  compasses 
sheathed  with  cork,  and  in  ascertaining  how  close  the  points  of  the  com- 
passes may  be  brought  to  each  other  and  still  be  felt  as  two  bodies. 


684  THE    SENSES 

TABLE  OF  VARIATIONS  IN  THE  TACTILE  SENSIBILITY  OF  THE  DIFFERENT  PARTS 

OF  THE  SKIN. 

The  measurement  indicates  the  least  distance  at  which  the  two  blunted  points  of  a  pair 
of  compasses  can  be  separately  distinguished  as  two.       (E.  H.  Weber.) 

Tip  of  tongue i  mm. 

Palmar  surface  of  third  phalanx  of  forefinger 2  mm. 

Palmar  surface  of  second  phalanges  of  fingers 4  mm. 

Red  surface  of  under-lip 4  mm. 

Tip  of  nose 6  mm. 

Middle  of  dorsum  of  tongue 8  mm. 

Palm  of  hand i  o  mm. 

Center  of  hard  palate 12  mm. 

Dorsal  surface  of  first  phalanges  of  fingers 14  mm. 

Back  of  hand 25  mm. 

Dorsum  of  foot  near  toes 37  mm. 

Gluteal  region. 37  mm. 

Sacral  region 37  mm. 

Upper  and  lower  parts  of  forearm 37  mm. 

Back  of  neck  near  occiput'. 50  mm. 

Upper  dorsal  and  mid-lumbar  regions 50  mm. 

Middle  part  of  forearm ,.62  mm. 

Middle  of  thigh 62  mm. 

Mid-cervical  region    62  mm. 

Mid-dorsal  region 62  mm. 

In  the  case  of  the  limbs,  before  the  points  are  recognized  as  two,  they 
have  to  be  separated  further  when  the  line  joining  them  is  in  the  long  axis  of 
the  limb  than  when  in  the  transverse  direction. 

According  to  Weber  the  mind  estimates  the  distance  between  two  points 
by  the  number  of  unexcited  nerve  endings  which  intervene  between  the  two 
points  touched.  It  would  appear  that  a  certain  number  of  intervening  un- 
excited nerve  endings  is  necessary  before  two  points  touched  can  be  recog- 
nized as  separate,  and  the  greater  this  number  the  more  clearly  are  the  points 
of  contact  distinguished  as  separate.  The  delicacy  of  the  sense  of  touch  may 
be  very  much  increased  by  practice.  A  familiar  illustration  occurs  in  the 
case  of  the  blind,  who,  by  constant  practice,  can  acquire  the  power  of  reading 
raised  letters  the  forms  of  which  are  almost,  if  not  quite,  undistinguishable  by 
the  sense  of  touch  to  an  ordinary  person. 

The  different  degrees  of  sensitiveness  possessed  by  different  parts  may 
give  rise  to  errors  of  judgment  in  estimating  the  distance  between  two  points 
where  the  skin  is  touched.  Thus,  if  blunted  points  of  a  pair  of  compasses 
(maintained  at  a  constant  distance  apart)  be  slowly  drawn  over  the  skin  of 
the  cheek  toward  the  lips,  it  is  almost  impossible  to  resist  the  conclusion  that 
the  distance  between  the  points  is  gradually  increasing.  When  they  reach 
the  lips  they  seem  to  be  considerably  farther  apart  than  on  the  cheek.  Thus, 


SENSE    OF    TEMPERATURE  685 

too,  our  estimate  of  the  size  of  a  cavity  in  a  tooth  is  usually  exaggerated  when 
based  upon  sensations  derived  from  the  tongue  alone.  Another  curious 
illusion  may  here  be  mentioned.  If  we  close  the  eyes,  and  place  a  small 
marble  or  pea  between  the  crossed  fore  and  middle  fingers,  we  seem  to  be 
touching  two  marbles,  figure  480.  This  illusion  is  due  to  an  error  of  judg- 
ment. The  marble  is  touched  by  two  surfaces  which,  under  ordinary  cir- 
cumstances, could  be  touched  only  by  two  separate  marbles,  hence  the  mind, 
taking  no  cognizance  of  the  fact  that  the  fingers  are  crossed,  forms  the  con- 
clusion that  two  sensations  are  due  to  two  marbles. 

Sense  of  Temperature. — The  whole  surface  of  the  body  is  more  or 
less  sensitive  to  differences  of  temperature.  The  sensation  of  heat  is  distinct 
from  that  of  touch,  hence  it  would  seem  reasonable  to  suppose  that  there  are 
special  nerves  and  nerve  endings  for  temperature.  At  any  rate  the  power  of 
discriminating  temperature  may  remain  unimpaired  when  the  sense  of  touch 
is  temporarily  in  abeyance.  Thus  if  the  ulnar  nerve  be  compressed  at  the 
elbow  till  the  sense  of  touch  is  very  much  dulled  in  the  fingers  which  it  sup- 
plies, the  sense  of  temperature  remains  quite  unaffected.  And  in  certain 
diseases  of  the  cord  the  sense  of  touch  may  be  impaired  in  a  part,  and  tem- 
perature remain  undisturbed,  or  the  converse. 


FIG.  421. — Diagram  of  a  Part  of  the  Hand,  Showing  Distribution  of  Sense  Spots;  for 
touch,  A ;  for  heat,  B;  and  for  cold,  C.  In  A  the  skin  is  sensitive  except  at  the  parts  marked 
with  black;  in  B  and  C,  the  intensity  of  the  shading  represents  the  relative  sensitiveness. 
(Goldscheider.) 


The  mapping  oi  the  surface  of  a  part  of  the  skin  with  reference  to  its 
sensibility  to  temperature  reveals  the  fact  that  there  are  definite  heat  and 
cold  spots.  Furthermore,  the  areas  do  not  coincide,  leading  us  to  conclude 
that  there  are  two  distinct  sense  organs  concerned,  figure  421,  B  and  C. 

The  sensations  of  heat  and  cold  are  often  exceedingly  fallacious,  and  in 
many  cases  are  no  guide  at  all  to  the  absolute  temperature  as  indicated  by 
a  thermometer.  All  that  we  can  with  safety  infer  from  our  sensations  of 
temperature  is  that  a  given  object  is  warmer  or  cooler  than  the  skin.  Thus 


686  THE   SENSES 

the  temperature  of  our  skin  is  the  standard;  and  as  this  varies  from  hour  to 
hour  according  to  the  activity  of  the  cutaneous  circulation,  our  estimate  of 
the  absolute  temperature  of  any  body  must  necessarily  vary  too.  If  we  put 
the  left  hand  into  water  at  5°  C.  (40°  F.)  and  the  right  into  water  at  45°  C. 
(110°  F.),  and  then  immerse  both -in  water  at  27°  C.  (80°  F.),  it  will  feel  warm 
to  the  left  hand,  but  cool  to  the  right.  Again,  a  piece  of  metal  which  has 
really  the  same  temperature  as  a  given  piece  of  wood  will  feel  much  colder, 
since  it  conducts  away  the  heat  much  more  rapidly.  For  the  same  reason 
air  in  motion  feels  very  much  cooler  than  air  of  the  same  temperature  at  rest. 

In  some  cases  we  are  able  to  form  a  fairly  accurate  estimate  of  absolute 
temperature.  Thus,  by  plunging  the  elbow  into  a  bath,  a  practiced  bath- 
attendant  can  tell  the  temperature  sometimes  within  half  a  degree 
centigrade. 

The  temperatures  which  can  be  readily  discriminated  are  between  10°  and 
45°  C-  (50°  and  115°  F.);  very  low  and  very  high  temperatures  alike  produce 
a  burning  sensation.  A  temperature  appears  higher  according  to  the  extent 
of  cutaneous  surface  exposed  to  it.  Thus,  water  of  a  temperature  which 
can  be  readily  borne  by  the  hand  is  quite  intolerable  if  the  whole  body  be 
immersed. 

The  delicacy  of  the  sense  of  temperature  coincides  in  the  main  with 
that  of  touch,  though  at  the  elbow  where  the  skin  is  thin,  and  the  sense  of 
temperature  is  delicate,  that  of  touch  is  not  remarkably  so.  Weber  has 
further  ascertained  two  points  so  near  together  on  the  skin  that  they  pro- 
duce but  a  single  impression,  at  once  give  rise  to  two  sensations  when  one 
is  hotter  than  the  other.  Moreover,  of  two  bodies  of  equal  weight,  that 
which  is  the  colder  feels  heavier  than  the  other. 

As  every  sensation  is  attended  with  a  perception  and  leaves  behind  it  an 
idea  in  the  mind  which  can  be  reproduced  at  will,  we  are  enabled  to  compare 
the  idea  of  a  past  sensation  with  another  sensation  really  present.  Thus  we 
can  compare  the  weight  of  one  body  with  another  which  we  had  previously 
felt,  of  which  the  idea  is  retained  in  our  mind.  Weber  was  indeed  able  to 
distinguish  in  this  manner  between  temperatures  experienced  one  after 
the  other,  better  than  between  temperatures  to  which  the  two  hands  were 
simultaneously  subjected.  This  power  of  comparing  present  with  past  sensa- 
tions diminishes,  however,  in  proportion  to  the  time  which  has  elapsed  between 
them.  After-sensations  left  by  impressions  on  nerves  of  common  sensibility 
or  touch  are  very  vivid  and  durable.  As  long  as  the  condition  into  which 
the  stimulus  has  thrown  the  organ  endures,  the  sensation  also  remains, 
though  the  exciting  cause  should  have  long  ceased  to  act.  Both  painful  and 
pleasurable  sensations  afford  many  examples  of  this  fact. 

Sense  of  Pain. — As  regards  painful  sensations,  three  views  can  be  taken, 
i,  That  it  is  a  special  sensation  provided  with  a  special  conducting  apparatus 
in  each  part  of  the  body;  2,  that  it  is  produced  by  an  over-stimulation  of  the 


THE    MUSCULAR   SENSE  687 

special  nerves  concerned  with  touch  or  temperature,  or  of  the  other  nerves 
of  special  sense;  or  3,  that  it  is  an  over-stimulation  of  the  nerves  of  common 
sensation,  which  tell  us  of  the  condition  of  our  bodies,  both  of  the  surface  and 
also  of  the  internal  organs.  There  seems  to  be  much  in  favor  of  all  of  these 
views.  The  weight  of  evidence  is,  however,  rather  against  there  being  any 
special  pain  sense  with  a  special  end-organ  and  fibers,  though  Barker  in  his 
own  arm  experienced  the  presence  of  pain  sensations  while  there  was  absence 
of  sensations  of  touch  and  temperature.  It  is,  indeed,  certain  that,  even  if 
any  variety  of  pain  be  a  special  sensation,  some  kind  of  pain  may  be  produced 
by  stimulation  of  the  bare  sensory  nerves  apart  from  any  special  form  of 
nerve  termination.  It  is  said  that  the  main  difference  between  the  common 
sensory  apparatus  which  tells  us  of  the  condition  of  all  parts  of  the  body  of 
which  thirst  and  hunger  are  but  examples,  and  the  special  sense  of  touch  and 
temperature,  is  that  the  latter  are  provided  with  a  special  local  apparatus. 
By  means  of  this  apparatus  we  are  able  to  localize  the  sensation.  Such  a 
special  apparatus  is  evidently  not  absolutely  essential  for  the  sensation  of 
pain,  but  this  does  not  exclude  the  idea  that  pain  may  result  from  over-stimu- 
lation of  a  nerve  of  special  sense  or  of  its  termination. 

The  Muscular  Sense. — The  estimate  of  a  weight  is  usually  based  on 
two  sensations:  i,  of  pressure  on  the  skin,  and  2,  the  sense  of  muscular 
resistance. 

The  estimate  of  weight  derived  from  a  combination  of  these  two  sensations 
(as  in  lifting  a  weight)  is  more  accurate  than  that  derived  from  the  former 
alone  (as  when  a  weight  is  laid  on  the  hand) ;  thus  Weber  found  that  by  the 
former  method  he  could  generally  distinguish  19^  oz.  from  20  oz.,  but  not 
19!  oz.  from  20,  while  by  the  latter  he  could  at  most  distinguish  only  14  J  oz. 
from  15  oz.  It  is  not  the  absolute,  but  the  relative,  amount  of  the  difference 
of  weight  which  we  have  thus  the  faculty  of  perceiving. 

It  is  not,  however,  certain,  that  our  idea  of  the  amount  of  muscular  force 
used  is  derived  solely  from  the  muscular  sense.  We  have  the  power  of  esti- 
mating very  accurately  beforehand,  and  of  regulating,  the  amount  of  nervous 
influence  necessary  for  the  production  of  a  certain  degree  of  movement. 
When  we  lift  a  vessel,  with  the  contents  of  which  we  are  not  acquainted, 
the  force  we  employ  is  determined  by  the  idea  we  have  conceived  of  its  weight. 
If  it  should  happen  to  contain  some  very  heavy  substance,  as  quicksilver, 
we  would  probably  fail  in  the  attempt;  the  amount  of  muscular  action,  or 
of  nervous  energy,  which  we  exerted  being  insufficient.  It  is  possible 
that  in  the  same  way  the  idea  of  weight  and  pressure  in  raising  bodies,  or  in 
resisting  forces,  may  in  part  arise  from  a  consciousness  of  the  amount  of 
nervous  energy  transmitted  from  the  brain  rather  than  from  a  sensation 
in  the  muscles  themselves.  The  mental  conviction  of  the  inability  longer  to 
support  a  weight  must  also  be  distinguished  from  the  actual  sensation  of 
fatigue  in  the  muscles. 


688  THE    SENSES 

So,  with  regard  to  the  ideas  derived  from  sensations  of  touch  combined 
with  movements,  it  is  doubtful  how  far  the  consciousness  of  the  extent  of 
muscular  movement  is  obtained  from  sensations  in  the  muscles  themselves. 
The  sensation  of  movement  attending  the  motions  of  the  hand  is  very  slight; 
and  persons  who  do  not  know  that  the  action  of  particular  muscles  is  neces- 
sary for  the  production  of  given  movements,  do  not  suspect  that  the  move- 
ment of  the  fingers,  for  example,  depends  on  an  action  in  the  forearm. 
The  mind  has,  nevertheless,  a  very  definite  knowledge  of  the  changes 
of  position  produced  by  movements;  and  it  is  on  this  that  the  ideas  which 
it  conceives  of  the  extension  and  form  of  a  body  are  in  great  measure 
founded. 

There  is  no  marked  development  of  common  sensibility  to  be  made  out 
in  muscles:  they  may  be  cut  without  the  production  of  pain.  On  the 
other  hand,  there  is  no  doubt  that  afferent  impulses  must  pass  upward 
from  muscles  and  tendons  to  the  brain  on  the  basis  of  which  we  become 
conscious  of  their  condition.  This,  then,  must  be  a  special  sense.  It  has 
been  suggested  that  the  minute  end-bulbs  of  Golgi  found  in  tendons,  and 
that  the  Pacinian  corpuscles  in  the  neighborhood  of  joints,  are  the  terminal 
organs  of  this  special  sense. 

Cutaneous  Sensibility  and  Differential  Innervation  of  the  Skin.— 
Studies  on  the  regeneration  of  cutaneous  or  sensory  nerve  trunks  after 
section  or  after  degeneration  from  disease  have  revealed  the  interesting 
fact  that  the  skin  possesses  two  types  of  sensory  nerves.  In  the  recovery 
of  sensibility  to  touch  and  temperature  and  to  pain  Head  and  Rivers  have 
discovered  that  after  a  relatively  short  time,  from  seven  to  twenty-six 
weeks,  a  certain  degree  of  sensitiveness  appears.  However,  the  sensations 
are  different  from  ordinary  sensibility — less  distinct,  more  diffuse,  different 
in  quality  and  cannot  be  localized  with  the  usual  definiteness.  The  stimu- 
lus to  evoke  them  must  be  coarser  and  more  general.  Only  relatively  wide 
extremes  of  heat  are  perceived,  heat  above  38°  C.  or  below  24°  C.  Pain 
sensations  are  more  disturbing,  of  a  peculiar  type.  This  type  of  sensibility 
has  been  designated  as  protopathic  to  distinguish  it  from  the  usual  type  or 
epicritic  sensibility.  In  regeneration  epicritic  sensibility  returns  much 
later,  in  two  years  more  or  less,  after  nerve  section  or  degeneration.  With 
the  return  of  the  epicritic  sensory  function  the  usual  accurate  discrimina- 
tion of  temperature  variations  within  the  narrower  limits  between  24° 
and  38°C.,  and  of  delicate  touch  sensibility,  accurate  spatial  localization, 
and  discrimination  replace  the  protopathic  type. 

These  experiments  lend  strength  to  the  view  that  cutaneous  sensibility 
depends  on  a  double  or  at  least  a  differential  innervation.  However, 
Head's  later  experiments  seem  to  show  that  the  two  classes  of  nerves 
probably  run  in  common  tracts  in  the  cord  and  brain  stem.  In  cord  lesions 
that  affect  the  epicritic  sensibility  the  protopathic  is  also  lost. 


TASTE    AND    SMELL  689 

Touch  Sensations  and  Judgment  of  the  Form  and  Size  of  Bodies. — By 

the  sense  of  touch  the  mind  is  made  acquainted  with  the  size,  form,  and 
other  external  characters  of  bodies.  And  in  order  that  these  characters 
may  be  easily  ascertained,  the  sense  of  touch  is  especially  developed  in 
those  parts  which  can  be  readily  moved  over  the  surface  of  bodies.  Touch, 
in  its  more  limited  sense,  or  the  act  of  examining  a  body  by  the  touch, 
consists  merely  in  a  voluntary  employment  of  this  sense  combined  with 
movement,  and  stands  in  the  same  relation  to  the  sense  of  touch,  or  com- 
mon sensibility,  generally,  as  the  act  of  seeking,  following,  or  examining 
odors  does  to  the  sense  of  smell.  The  hand  is  the  best  adapted  for  it,  by 
teason  of  its  peculiarities  of  structure — namely,  its  capability  of  pronation 
and  supination,  which  enables  it,  by  the  movement  of  rotation,  to  examine 
rhe  whole  circumference  of  a  body;  the  power  it  possesses  of  opposing  the 
thumb  to  the  rest  of  the  hand,  and  the  relative  mobility  of  the  ringers; 
and  lastly  from  the  abundance  of  the  sensory  terminal  organs  which  it 
possesses.  In  forming  a  conception  of  the  figure  and  extent  of  a  surface, 
the  mind  multiplies  the  size  of  the  hand  or  fingers  used  in  the  inquiry  by 
the  number  of  times  which  it  is  contained  in  the  surface  traversed;  and, 
by  repeating  this  process  with  regard  to  the  different  dimensions  of  a  solid 
body,  acquires  a  notion  of  its  cubical  extent,  but,  of  course,  only  an  imper- 
fect notion,  as  other  senses,  e.g.,  the  sight,  are  required  to  make  it  complete. 
It  is  impossible  in  this  consideration  to  say  how  much  of  our  knowledge 
of  the  thing  touched  depends  upon  pressure  and  how  much  upon  the  mus- 
cular sense. 

II.  TASTE  AND  SMELL. 

The  special  sense  organs  for  taste  and  smell  are  stimulated  by  chemical 
substances,  the  former  by  chemicals  in  solution,  the  latter  by  volatile  materials. 
They  are  also  closely  associated  in  action  and  we  do  not  always  differentiate 
between  the  two. 

THE  SENSE  OF  TASTE. 

The  conditions  for  the  perceptions  of  taste  are:  i,  the  presence  of  a  sense 
organ,  a  nerve,  and  a  nerve  center  with  special  endowments;  2,  the  excitation 
of  the  sense  organ  by  the  sapid  matters,  which  for  this  purpose  must  be  in  a 
state  of  solution;  3,  a  temperature  of  about  37°  to  40°  C.  (98°  to  100°  F.). 

The  Nerves  and  Organs  of  Taste. — The  principal  organ  of  the  sense 
of  taste  is  the  tongue.  But  the  soft  palate  and  its  arches,  the  uvula,  tonsils, 
and  probably  the  upper  part  of  the  pharynx,  are  also  endowed  with  taste. 
These  parts,  together  with  the  base  and  posterior  parts  of  the  tongue,  are 
supplied  with  branches  of  the  glosso-pharyngeal  nerve,  and  evidence  has 

44 


THE    SENSES 


been  already  adduced  that  this  is  the  principal  nerve  of  the  sense  of  taste. 
The  anterior  parts  of  the  tongue,  especially  the  edges  and  tip,  are  innervated 
by  fibers  from  the  lingual  branch  of  the  fifth,  but  which  arise  in  the  ganglion 
of  the  pars  intermedia  and  are  distributed  in  the  chorda  tympani,  figures  250 
and  412. 


FIG.  422. — Papillar  Surface  of  the  Tongue,  with  the  Fauces  and  Tonsils,  i,  Circum- 
vallate  papillae,  in  front  of  2,  the  foramen  cecum;  3,  fungiform  papillae;  4,  filiform  and 
conical  papillae;  5,  transverse  and  oblique  rugae;  6,  mucous  glands  at  the  base  of  the  tongue 
and  in  the  fauces;  7,  tonsils;  8,  part  of  the  epiglottis;  9,  median  glosso-epiglottidean  fold 
(frenum  epiglottidis).  (From  Sappey.) 

The  mucous  membrane  in  the  regions  just  mentioned  possesses  special 
epithelial  structures  called  taste  buds.  The  taste  buds  are  very  abundant 
in  the  side  walls  of  the  circumvallate  papillae.  They  are  also  present  in  the 
fungiform  papillae,  in  the  foliate  papillae,  and  in  the  mucous  membrane. 
The  taste  bud  is  located  at  the  deeper  part  of  the  stratified  epithelium,  is 


THE    SENSE    OF    TASTE 


691 


ovoid  in  shape,  and  its  free  end  abuts  on  the  surface  or  opens  to  the  surface 
by  a  short  canal.  It  is  composed  of  two  kinds  of  modified  epithelial  cells — 
the  supporting  cells,  which  are  long,  spindle-shaped  cells  that  form  a  sheath 
around  the  special  gustatory  cells;  and  the  taste  cells,  which  are  neuro-epithe- 
lial  cells  that  are  found  in  the  center  of  the  taste  bud.  They  are  very  slender 
cells  that  project  on  the  surface  by  a  delicate  process.  A  bundle  of  nerve 
fibrils  enters  the  base  of  each  taste  bud  and  forms  a  net  about  the  taste  cells. 


FIG.  423. 


FIG.  424. 


FIG.  423. — Taste  Bud  from  Side  Wall  of  Circumvallate  Papillae.  (Merkel-Henle.) 
a,  Taste  pore;  b,  nerve  fibers,  some  of  which  enter  the  taste  bud,  intrageminal  fibers,  while 
others  end  freely  in  the  surrounding  epithelium,  intergeminal  fibers. 

FIG.  424. — Vertical  Section  of  a  Circumvallate  Papilla  of  the  Calf,  i  and  3,  Epithelial 
layers  covering  it;  2,  taste  goblets;  4,  and  4',  duct  of  serous  gland  opening  out  into  the  pit 
in  which  the  papilla  is  situated;  5  and  6,  nerves  ramifying  within  the  papilla.  (Engelmann.) 

The  Circumvallate,  the  fungiform,  and  the  filiform  papillae,  shown  in 
figure  422,  are  special  structures  that  facilitate  the  stimulation  of  the  taste 
buds  by  sapid  substances.  They  are  all  formed  by  a  projection  of  the 
mucous  membrane,  and  contain  special  branches  of  blood  vessels  and 
nerves.  In  details  of  structure,  however,  they  differ  considerably  one  from 
another. 

Circumvallate  Papilla. — These  papillae,  figure  424,  eight  or  ten  in  nu'mber, 
are  situated  in  two  V-shaped  lines  on  the  base  of  the  tongue.  They  are 
circular  elevations  from  i  to  2  mm.  in  diameter  each,  with  a  central  depres- 
sion, and  surrounded  by  a  circular  fissure,  at  the  outside  of  which  is  a  slightly 
elevated  ring.  Both  the  central  elevation  and  the  ring  are  formed  of  close 
set  simple  papillae. 

Fungiform  Papilla. — The  fungiform  papillae  are  scattered  chiefly  over 
the  sides  and  tip,  and  sparingly  over  the  middle  of  the  dorsum,  of  the  tongue; 
the  name  is  derived  from  their  being  usually  narrower  at  the  base  than  at 
the  summit.  They  also  are  supplied  with  loops  of  capillary  blood  vessels, 
and  nerve  fibers. 


g2  THE    SENSES 

Conical  or  Filiform  Papilla. -^These,  which  are  the  most  abundant 
papillae,  are  scattered  over  the  whole  surface  of  the  tongue,  but  especially 
over  the  middle  of  the  dorsum.  They  vary  in  shape  somewhat,  but  for  the 
most  part  are  conical. 

Taste  Sensations. — The  occurrence  of  two  kinds  of  special  sensi- 
bility, i.e.,  touch  and  taste  in  the  same  part,  makes  it  sometimes  difficult  to 
determine  whether  the  impression  produced  by  a  substance  is  perceived 
through  the  ordinary  tactile  sensitive  fibers,  or  through  those  of  the  sense  of 
taste.  In  many  cases,  indeed,  it  is  probable  that  both  sets  of  nerve  fibers 
are  concerned,  as  when  irritating  acrid  substances  are  introduced  into  the 
mouth. 

Many  of  the  so-called  tastes  are  due  to  the  sapid  substances  being  also 
odorous,  and  exciting  the  simultaneous  action  of  the  sense  of  smell.  This  is 
shown  by  the  insipid  taste  of  certain  substances  when  their  action  on  the 
olfactory  nerves  is  prevented  by  closing  the  nostrils.  Many  of  the  popular 
drinks  lose  much  of  their  apparent  excellence  if  the  nostrils  are  held  close 
while  they  are  drunk. 

When  these  accessory  sensations  are  taken  into  account  it  is  found  that 
the  clearly  defined  tastes  are  reduced  to  four:  sweet,  bitter,  acid,  and  salt. 
These  taste  sensations  are  produced  by  the  respective  substances  when  in 
solution.  If  dry  salt  or  quinine  is  placed  on  the  surface  of  the  tongue,  no 
taste  appears  until  solution  takes  place  in  the  secretions  of  the  tongue.  A 
piece  of  metal,  as  a  silver  coin,  gives  rise  to  a  seemingly  distinct  taste  sensa- 
tion, called  metallic,  but  it  is  probably  not  to  be  accepted  as  co-ordinate  with 
the  others.  The  acid  taste  may  be  excited  by  electricity.  If  a  piece  of  zinc 
be  placed  beneath  and  a  piece  of  copper  above  the  tongue,  and  their  ends 
brought  into  contact,  an  acid  taste  (due  to  the  feeble  galvanic  current)  is 
produced.  The  delicacy  of  the  sense  of  taste  is  sufficient  to  discern  one  part 
of  sulphuric  acid  in  10,000  of  water,  or  one  part  of  quinine  in  200,000  of 
water.  But  it  is  far  surpassed  in  acuteness  by  the  sense  of  smell. 

ACUTENESS  OF  THE  SENSE  OF  TASTE.        (HALL.) 

The  average  of  10  individuals. 

Sugar i  part  to  520 

Quinine i  part  to  444,000 

Acetic  acid i  part  to        5,640 

Salt i  part  to          469 

Exploration  of  the  taste  areas  reveals  the  fact  that  regions  of  the  tongue 
and  mouth  are  not  equally  sensitive  to  the  sapid  substances.  Sweet  tastes 
are  especially  developed  at  the  tip  and  sides  of  the  tongue,  while  bitter  tastes 
are  almost  absent  in  the  front,  but  especially  developed  on  the  basal  region, 
and  in  the  fauces  and  pharynx.  Salts  are  more  stimulating  to  the  tip  of  the 
tongue,  and  acids  along  the  sides.  Individual  tests  of  the  fungiform  papillae 


AFTER- TASTES  AND  CONTRASTS 


693 


by  Oehrwall  showed  that  about  half  the  papillae  reacted  to  sweet,  bitter,  and 
acid,  but  that  certain  ones  reacted  only  to  sweet,  or  to  sweet  and  bitter,  or 
to  acid  and  bitter.  This  suggests  the  specific  nature  of  the  taste  sensations 
and  tends  to  prove  that  there  may  be  a  special  organ  for  each  kind  of  stimulus. 
Experiments  have  also  shown  that  it  is  possible  to  do  away  with  the  power 
of  tasting  bitters  and  sweets  while  the  taste  for  acids  and  salts  remains. 
This  is  done  by  chewing  the  leaves  of  an  Indian  plant,  Gymnema  sylvestre. 
Jt  has  also  been  shown  that  the  power  of  tasting  sweet  substances  disappears 
before  that  of  tasting  bitter.  Other  experiments  have  shown  that  the  mech- 
anisms for  salt  and  acid  tastes  are  distinct. 


FIG.  425. — Localization  of  Taste.     Bitter ;   acid ;   salt, — . — . — ;  sweet ;  T, 

tonsils;  FC,  foramen  cecum;  CF,  circumvallate  papillae;  FP,  fungiform  papillae.     (Hall.) 


After-tastes  and  Contrasts. — Very  distinct  sensations  of  taste  are 
frequently  left  after  the  substances  which  excited  them  have  ceased  to  act 
on  the  nerve,  as  the  after- taste  of  metallic  bitter,  which  remains  after  breaking 
the  stimulating  current.  Such  sensations  often  endure  for  a  long  time,  and 
modify  the  taste  of  other  substances  applied  to  the  tongue.  Thus,  the  taste 
of  sweet  substances  is  intensified  after  the  tasting  of  common  salt.  After 
rinsing  the  mouth  with  water  containing  salt,  it  is  said  that  sweet  solutions 
are  perceived  that  are  too  dilute  to  be  detected  ordinarily.  Many  other 
chemicals  produce  similar  results.  The  application  of  a  sapid  substance, 
acid  for  example,  to  one  side  of  the  tongue  intensifies  the  sensation  produced 
by  a  sapid  substance  applied  to  the  other  side.  There  is  a  simultaneous  con- 
trast which  suggests  that  the  same  relation  exists  between  tastes  as  between 
colors,  of  which  those  that  are  opposed,  i.e.,  complementary,  render  each 
other  more  vivid,  though  no  general  principles  governing  this  relation  have 


694  THE    SENSES 

been  discovered  in  the  case  of  tastes.  In  the  art  of  cooking,  however,  atten- 
tion has  at  all  times  been  paid  to  the  consonance  or  harmony  of  flavors  in 
their  combination  or  order  of  succession,  just  as  in  painting  and  music  the 
fundamental  principles  of  harmony  have  been  employed  empirically  while 
the  theoretical  laws  were  unknown. 

Frequent  and  continued  repetitions  of  the  same  taste  render  the  perception 
of  it  less  and  less  distinct,  in  the  same  way  that  a  color  becomes  more  and 
more  dull  and  indistinct  the  longer  the  eye  is  fixed  upon  it.  There  is  fatigue 
of  the  taste  organ  at  some  point. 

THE  SENSE  OF  SMELL. 

The  sensation  of  smell  is  produced  by  the  action  of  odorous  particles  on  a 
special  end-apparatus,  which  in  turn  causes  nerve  impulses  that  arouse 
changes  in  a  special  area  in  the  sensorium.  The  stimulating  cause  is  the 
direct  action  of  chemical  substances  as  in  the  sense  of  taste.  In  this  case 


FIG.  426. — Nerves  of  the  Septum  Nasi,  Seen  from  the  Right  Side.  X  f. — /,  The 
olfactory  bulb;  i,  the  olfactory  nerves  passing  through  the  foramina  of  the  cribriform  plate, 
and  descending  to  be  distributed  on  the  septum;  2,  the  internal  or  septal  twig  of  the  nasal 
branch  of  the  ophthalmic  nerve;  3,  naso-palatine  nerves.  (From  Sappey,  after  Hirschfeld 
and  Leveille.) 


however,  the  substances  must  reach  the  sensory  membrane  in  a  gaseous 
state  or  in  extremely  fine  division,  so  that  it  can  quickly  enter  into  solution 
in  the  moisture  on  the  sensitive  mucous  surface.  The  odorous  particles  are 
carried  to  the  membrane  by  inspiratory  currents  of  air. 

The  Olfactory  Apparatus. — The  essential  parts  of  the  olfactory  ap- 
paratus are  the  nasal  sensory  or  olfactory  membrane  to  receive  the  special 
stimuli,  and  the  nervous  apparatus  to  conduct  the  olfactory  nerve-impulse  to 
the  sensory  area  in  the  cortex  cerebri  for  its  perception. 


THE  OLFACTORY  APPARATUS  695 

The  nose  is  not  entirely  an  organ  for  the  seat  of  smell.  In  fact  the  nasal 
cavities  are  divided  into  three  districts  called,  respectively :  i ,  Regio  vestibularis 
which  is  the  entrance  to  the  cavity.  It  is  lined  with  a  mucous  membrane  very 
closely  resembling  the  skin,  and  guarded  by  hairs  and  by  sebaceous  glands. 
2,  Regio  respiratorid,  which  includes  the  lower  and  middle  meatus  of  the 
nose.  It  is  covered  with  mucous  membrane  of  stratified  columnar  ciliated 
epithelium.  The  mucosa  is  thick  and  consists  of  fibrous  connective  tissue; 
it  contains  a  certain  number  of  tubular  mucous  and  serous  glands.  3,  Regio 


FIG.  427. — Nerves  of  the  Outer  Walls  of  the  Nasal  Fossae,  i,  Network  of  the  branches 
of  the  olfactory  nerve,  descending  upon  the  region  of  the  superior  and  middle  turbinated 
bones;  2,  external  twig  of  the  ethmoidal  branch  of  the  nasal  nerves;  3,  spheno-palatine 
ganglion;  4,  ramification  of  the  anterior  palatine  nerves;  5,  posterior,  and  6,  middle  divisions 
of  the  palatine  nerves;  7,  branch  to  the  region  of  the  inferior  turbinated  bone;  8,  branch  to 
the  region  of  the  superior  and  middle  turbinated  bones;  9,  naso-palatine  branch  to  the 
septum  cut  short.  (From  Sappey,  after  Hirschfeld  and  Leveille.) 

olfactoria.  This  includes  the  anterior  two-thirds  of  the  superior  meatus,  the 
middle  meatus,  and  the  upper  half  of  the  septum  nasi,  figures  427  and  428. 
It  is  of  a  yellowish  color.  It  consists  of  a  thicker  mucous  membrane  than 
in  2,  made  up  of  loose,  areolar  connective  tissue  covered  by  epithelium  of  a 
special  variety,  resting  upon  a  basement  membrane.  The  cells  of  the  epithe- 
lium are  of  two  principal  kinds:  0,  columnar  epithelial  cells  whose  function 
is  to  support  b,  the  bipolar  olfactory  cells.  The  epithelial  cells  are  prismatic 
in  shape  and  have  upon  their  surfaces  facets  into  which  the  olfactory  cells 
fit  themselves,  figure  428,  e.  They  are  thus  analogous  to  the  cells  of  Miiller 
of  the  retina.  The  olfactory  cells  have  an  oblong  or  fusiform  shape,  which 
is  mainly  determined  by  the  large  nucleus.  The  thin  protoplasmic  body  has 
two  processes,  an  external  and  an  internal.  The  external  is  large  and  passes 
up  to  the  free  surface,  to  end  in  a  small  bunch  of  fibrils  that  are  not  vibratile. 
The  internal  process  is  very  fine,  often  varicose,  and  passes  through  the 


THE    SENSES 


cribriform  plate  to  form  a  glomerular  basket  with  the  branches  of  the  mitral 
cells  of  the  olfactory  bulb. 

The  olfactory  bulb  must  be  studied  in  relation  with  the  nerve  fibers  and 
olfactory  cells  with  which  it  is  connected.  These  parts  together  form  a  sen- 
sory end-organ  which  resembles  in  many  respects  the  retina.  The  discovery 

of  its  true  structure  has  thrown  a  flood  of  light  on 
the  architecture  of  the  nerve  centers  as  a  whole. 

The  olfactory  bulb  is  not  a  nerve,  but  a  modi- 
fication of  the  brain  cortex.  A  transection  shows 
it  to  be  made  up  of  four  layers:  i.  Peripheral 
fibers.  2.  Olfactory  glomerules.  3.  Layer  of 
mitral  cells.  4.  Layer  of  granular  cells  and  deep 
nerve  fibers. 

The  first  and  external  layer  is  composed  of 
the  fine  nerve  fibrils  of  the  olfactory  nerves. 
They  pass  through  the  cribriform  plate  of  the 
ethmoid,  arising  from  the  olfactory  cells  of  which 
they  are  processes. 

The  glomerular  layer  contains  numbers  of 
small  round  bodies  whose  structure  shows  that 
they  are  made  up  of  the  interlocking  expansions 
of  the  olfactory  fibers,  on  the  one  hand,  and  of 
the  branches  of  the  " mitral"  cells,  on  the  other. 
These  are  mingled  in  a  close  network,  but  do  not 
anastomose.  It  was  by  the  study  of  these  bodies 
in  part  that  the  fact  of  the  non-continuity  of  the 
neurones  was  demonstrated,  figure  429.  This 
layer  also  contains  small  fusiform  cells  with 
branching  dendrites  that  extend  outward  to  the 
glomeruli.  Each  has  an  axis-cylinder  process 
which  passes  inward  to  join  the  fibers  of  the 
internal  olfactory  nerves. 

The  layer  of  mitral  cells  contains  large  cells, 
some  of  them  triangular  and  some  in  the  shape  of  a  miter.  They  have 
numerous  dendrites,  one  of  which  passes  into  a  glomerulus  and  then  breaks 
up  in  a  fine  arborization.  An  axis-cylinder  process  passes  off  from  the 
inner  surface  and  is  continued  as  an  internal  olfactory  nerve  fiber  in  the 
olfactory  tract. 

The  layer  of  granules  and  central  fibers  contains  a  large  number  of  very 
small  nerve  cells,  which  are  peculiar  in  that  they  have  no  axis- cylinder. 
Their  dendrites  extend  chiefly  into  the  layer  of  mitral  cells.  They  resemble 
the  spongioblasts  of  the  retina  and  probably  have  commissural  functions. 
This  layer  has  also  some  small  star-shaped  cells  whose  dendrites  end  in  the 


FIG.  428.— Bipolar  Olfac- 
tory Cells  from  the  Nasal 
Fossae  of  the  Rat  (Full-term 
Fetus).  Ay  Epithelium  of  the 
olfactory  mucosa;  ey  epithelial 
cells;/,/,  nerve  cells;  i,  nerve 
fibers  terminating  freely  on 
the  epithelial  surface;  h, 
olfactory  nerve  fibers;  g,  sen- 
sory nerve  derived  from  the 
trigeminus.  (Cajal.) 


THE    STIMULATION    OF    THE    OLFACTORY    MEMBRANE 


697 


mitral-cell  layer.  Among  these  cells  run  numerous  fibers,  chiefly  from  the 
mitral  cells  and  the  fusiform  cells  of  the  glomerular  layer.  The  general 
arrangement  is  shown  in  figure  429. 

The  Stimulation  of  the  Olfactory  Membrane. — The  extent  of  the 
nasal  mucous  surfaces,  and  of  the  frontal  and  antral  sinuses  connected  with 
them,  might  suggest  that  the  sensory  olfactory  surface  is  widely  distributed, 
but  such  is  not  the  case.  Air  impregnated  with  vapor  of  camphor  has  been 
injected  into  the  frontal  sinus  through  a  fistulous  opening,  and  odorous  sub- 
stances have  been  injected  into  the  antrum  of  Highmore;  but  in  neither  case 


FIG.  429. — Principal  Constituent  Elements  of  the  Olfactory  Bulb  of  a  Mammal. 

Gehuchten.) 


(Van 


was  any  odor  perceived  by  the  patient.  All  parts  of  the  nasal  cavities  are 
endowed  with  cutaneous  sensibility  by  the  nasal  branches  of  the  first  and 
second  divisions  of  the  fifth  nerve,  hence  the  sensations  of  cold,  heat,  itching, 
tickling,  and  pain,  and  the  sensation  of  tension  or  pressure  in  the  nostrils. 
That  these  nerves  cannot  perform  the  functions  of  the  olfactory  nerves  is 
proved  by  cases  in  which  the  sense  of  smell  is  lost,  while  the  mucous  mem- 
brane of  the  nose  remains  susceptible  to  the  various  modifications  of  the 
sense  of  touch.  But  it  is  often  difficult  to  distinguish  the  sensation  of  smell 
from  that  of  mere  feeling,  and  to  ascertain  what  belongs  to  each  separately. 
This  is  true  particularly  of  the  sensations  excited  by  acrid  vapors  in  the  nose, 


698  THE    SENSES 

as  of  ammonia,  horse-radish,  mustard,  etc.,  and  the  difficulty  is  the  greater 
when  it  is  remembered  that  these  acrid  vapors  have  nearly  the  same  action 
upon  the  mucous  membrane  of  the  eyelids. 

The  true  olfactory  membrane  is  limited  to  the  small  area  on  either  side 
of  the  superior  meatus  and  supplied  by  the  olfactory  nerve.  It  is  stimulated 
by  odorous  substances  when  they  penetrate  the  upper  chamber  of  the  nose. 
Currents  of  air  can  be  drawn  over  this  membrane  more  certainly  and  effect- 
ively by  sniffing  the  air,  as  noticed  in  the  acts  of  a  dog  following  the  trail. 
The  odorous  particles  must  come  into  contact  with  the  olfactory  cells  when 
in  solution  in  the  moisture  over  the  surface  and  produce  the  stimulus  by  chem- 
ical change.  Mere  presence  in  solution  is  not  always  adequate  to  a  stimula- 
tion. It  seems  that  movement  over  the  surface  is  necessary,  at  least  to  effect- 
ive stimulation.  Haycraft  has  repeated  some  of  the  older  experiments  and 
finds  that  eau  de  Cologne  can  be  introduced  into  the  nasal  cavity  in  warm 
saline  solutions  without  producing  a  sensation  of  smell  even  when  10  per  cent, 
solutions  are  used.  He  also  showed  that  Cologne,  bergamot,  etc.,  can  be 
slowly  diffused  into  the  nasal  cavity  without  producing  a  stimulus.  If, 
while  the  vapor  is  thus  in  the  nasal  cavity,  the  nostril  be  closed  and  the 
person  goes  into  pure  air  and  breathes,  then  an  odorous  sensation  is  at  once 
experienced.  This  shows  that  even  odorous  gases  "must  be  moved  over  the 
olfactory  surface"  in  order  to  produce  a  stimulus. 

The  presence  of  bodies  in  quantities  so  minute  as  to  be  undiscernible 
even  by  spectrum  analysis,  0.000,000,03  of  a  grain  of  musk,  can  be  distinctly 
smelt  (Valentin).  Opposed  to  the  sensation  of  an  agreeable  odor  is  that  of 
a  disagreeable  or  disgusting  odor,  which  corresponds  to  the  sensations  of 
pain,  dazzling  and  disharmony  of  colors,  and  dissonance  in  the  other  senses. 
The»cause  01  this  difference  in  the  effect  of  different  odors  is  unknown;  but  this 
much  is  certain,  that  odors  are  pleasant  or  offensive  in  a  relative  sense  only, 
for  many  animals  pass  their  existence  in  the  midst  of  odors  which  to  us  are 
highly  disagreeable.  A  great  difference  in  this  respect  is,  indeed,  observed 
among  men.  Many  odors,  generally  thought  agreeable,  are  to  some  per- 
sons intolerable;  and  different  persons  describe  differently  the  sensations 
that  they  severally  derive  from  the  same  odorous  substances.  There  seems 
also  to  be  in  some  persons  an  insensibility  to  certain  odors,  comparable  with 
that  of  the  eye  to  certain  colors;  and  among  different  persons,  as  great  a 
difference  in  the  acuteness  of  the  sense  of  smell  as  among  others  in  the  acute- 
ness  of  sight.  We  have  no  exact  proof  that  a  relation  of  harmony  exists 
between  odors  as  between  colors  and  sounds,  though  it  is  probable  that  such 
is  the  case,  since  it  certainly  is  so  with  regard  to  the  sense  of  taste.  Such  a 
relation  would  account  in  some  measure  for  the  different  degrees  of  perceptive 
power  in  different  persons;  for  as  some  have  no  ear  for  music,  so  others  have 
no  clear  appreciation  of  the  relation  of  odors,  and  therefore  little  pleasure 
in  them. 


THE   EXTERNAL    EAR  699 

Most  of  the  substances  taken  as  foods  into  the  mouth  give  off  odorous 
particles  that  stimulate  the  olfactory  membrane.  In  fact,  the  chief  elements 
in  food  flavors  are  not  tastes,  but  smells,  or  combinations  of  the  two.  This 
is  particularly  true  of  meats.  Meats  are  especially  prized  for  their  delicate 
flavors,  and  cooking  is  performed  to  bring  out  these  flavors.  Yet  meat  has 
little  taste  other  than  salt;  the  so-called  tastes  are  due  to  odorous  particles 
entering  the  nostril  and  stimulating  the  olfactory  membrane  at  the  same 
moment  the  taste  buds  of  the  mouth  are  stimulated. 

Subjective  sensations  occur  frequently  in  connection  with  the  sense  of 
smell.  Often  a  person  smells  something  which  is  not  present,  and  which 
other  persons  cannot  smell;  this  is  very  frequent  with  nervous  persons,  but 
it  occasionally  happens  to  every  one.  In  a  man  who  was  conscious  of  a  bad 
odor,  the  arachnoid  was  found  after  death  to  be  beset  with  deposits  of  bone, 
and  a  lesion  in  the  middle  of  the  cerebral  hemispheres  was  also  discovered. 
Dubois  was  acquainted  with  a  man  who,  ever  after  a  fall  from  his  horse, 
which  occurred  several  years  before  his  death,  believed  that  he  smelt  a  bad 
odor. 

HEARING  AND  EQUILIBRATION. 
THE  ANATOMY  OF  THE  EAR. 

For  descriptive  purposes,  the  ear,  or  organ  of  hearing,  is  divided  into 
three  parts,  i,  the  external,  2,  the  middle,  and  3,  the  internal  ear.  The  first 
two  are  only  accessory  structures  to  the  third,  which  contains  the  essential 
parts  of  the  organ  of  hearing.  The  accompanying  figure,  430,  shows  very 
well  the  relation  of  these  divisions  to  each  other. 

The  External  Ear. — The  external  ear  consists  of  the  pinna  or  auricle 
and  the  external  auditory  canal  or  meatus. 

The  principal  parts  of  the  pinna,  figure  430,  are  two  prominent  rims  en- 
closed one  within  the  other,  the  helix  and  antiheliXj  and  inclosing  a  central 
hollow  named  the  concha;  in  front  of  the  concha,  a  prominence  directed 
backward,  the  tragus,  and  opposite  to  this  one  directed  forward,  the  anti- 
tragus.  From  the  concha,  the  auditory  canal  passes  inward  and  a  little 
forward  to  the  membrana  tympani,  to  which  it  thus  serves  to  convey  the 
vibrations  of  the  air.  It  consists  of  a  nbro-cartilage  tube  lined  by  skin  con- 
tinuous with  that  of  the  pinna,  and  extending  over  the  outer  part  of  the  mem- 
brana tympani.  Fine  hairs  and  sebaceous  glands  are  present  toward  the 
outer  part  of  the  canal,  while  deeper  in  the  canal  are  small  glands,  resembling 
the  sweat  glands  in  structure,  which  secrete  the  cerumen. 

Regarding  the  external  ear,  therefore,  as  a  collector  and  conductor  of 
sonorous  vibrations,  all  its  inequalities,  elevations,  and  depressions  become 
of  evident  importance;  for  those  elevations  and  depressions  upon  which  the 
undulations  fall  will  tend  to  intensify  certain  sound  waves  while  not  affecting 


700 


THE   SENSES 


FIG,  430. — Diagrammatic  View  from  Before  of  the  Parts  Composing  the  Organ  of 
Hearing  of  the  Left  Side.  The  temporal  bone  of  the  left  side,  with  the  accompanying 
soft  parts,  has  been  detached  from  the  head,  and  a  section  has  been  carried  through  it 
transversely,  so  as  to  remove  the  front  of  the  meatus  externus,  half  the  tympanic  membrane, 
the  upper  and  anterior  wall  of  the  tympanum  and  Eustachian  tube.  The  meatus  internus 
has  also  been  opened,  and  the  bony  labyrinth  exposed  by  the  removal  of  the  surrounding 
parts  of  the  petrous  bone,  i,  The  pinna  and  lobe;  2,  2',  meatus  externus;  2',  membrana 
tympani;  3,  cavity  of  the  tympanum;  3',  its  opening  backward  into  the  mastoid  cells; 
between  3  and  3',  the  chain  of  small  bones;  4,  Eustachian  tube;  5,  meatus  internus,  contain- 
ing the  facial  (uppermost)  and  the  auditory  nerves;  6,  placed  on  the  vestibule  of  the  laby- 
rinth above,  the  fenestra  ovalis;  a,  apex  of  the  petrous  bone;  6,  internal  carotid  artery;  c, 
styloid  process;  d,  facial  nerve  issuing  from  the  stylo-mastoid  foramen;  e,  mastoid  process; 
at  squamous  part  of  the  bone  covered  by  integument,  etc.  (Arnold.) 


s  B 


15     D 


FIG.  431. — Tympanic  Ossicles  of  Left  Ear.  A,  Incus  seen  from  the  front;  B,  malleus, 
viewed  from  behind;  C,  incus,  and  D,  malleus,  seen  from  inner  aspect;  E,  stapes,  i,  Body 
of  incus,  with  articular  surface  for  head  of  malleus;  2,  processus  longus;  3,  processus 
lenticularis;  4,  articular  surface  for  incus;  5,  head,  6,  neck;  7,  processus  brevis;  8,  manu- 
brium;  9,  body;  10,  short  process;  u,  long  process;  12,  processus  longus;  13,  head,  14, 
facet  for  incus;  15,  manubrium;  16,  head;  17,  neck;  18,  crus  anterius;  19,  crus  posteriusj 
20,  foot  plate. 


THE    MIDDLE    EAR    OR    TYMPANUM 


701 


others.  It  is  thought  that  this  forms  at  least  an  aid  in  determining  the 
direction  whence  a  sound  comes. 

The  Middle  Ear  or  Tympanum. — The  middle  ear,  or  tympanum,  3, 
figure  430,  is  separated  by  the  membrana  tympani  from  the  external  auditory 
canal.  It  is  a  cavity  in  the  temporal  bone,  opening  through  its  anterior  and 
inner  wall  into  the  Eustachian  tube. 

The  Eustachian  canal  establishes  communication  between  the  tympanic 
cavity  and  pharynx,  thus  equalizing  the  air  pressure  on  the  sides  of  the 
tympanic  membrane,  serving  the  same  mechanical  purpose  as  the  vent-hole 
in  a  snare  or  bass  drum.  The  cavity  of  the  tympanum  communicates  pos- 
teriorly with  air  cavities,  the  mastoid  cells,  in  the  mastoid  process  of  the  tem- 


Eecessus  epitympanicus 
Body  of  incus 

Short  process  of  incus 
Ligament  of  infc" 


Chorda  tympani  nerve 

Pyramid,  with  tendon 

of  stapedius  muscle' 

issuing  from  it 


Foot  of  stape 


Superior'Iigament  of  malleus 
•Head  of  malleus 

Anterior  ligament  of  malleus 
•Handle  of  malleus 

ffe 


Tensor  tympani  muscle 

rocessus 
ichleariformis 
Osseous  part  of 
Eustachian  tube 


FIG.  432. — Left  Membrana  Tympani  and  Chain  of  Tympanic  Ossicles  (Seen  from  Inner 

Aspect).     (Cunningham.) 

poral  bone;  but  its  only  opening  to  the  external  air  is  through  the  Eustachian 
tube.  The  cavity  of  the  tympanum  is  lined  with  mucous  membrane,  the 
epithelium  of  which  is  ciliated  and  continuous  with  that  of  the  pharynx.  It 
contains  a  chain  of  small  bones,  ossicula  auditus,  which  extends  from  the 
membrana  tympani  to  the  fenestra  ovalis. 

The  Membrana  Tympani. — The  tympanic  membrane  is  placed  in  a  slant- 
ing direction  at  the  bottom  of  the  external  canal,  its  plane  being  at  an  angle 
of  about  forty-five  degrees  with  the  lower  wall  of  the  canal.  It  is  formed 
chiefly  of  a  tough  and  tense  fibrous  membrane,  the  edges  of  which  are  set 
in  a  bony  groove.  Its  outer  surface  is  covered  by  a  continuation  of  the 
epithelial  lining  of  the  auditory  canal,  its  inner  surface  with  part  of  the 
mucous  membrane  of  the  middle  ear. 

The  Tympanic  Ossicles. — The  ear  bones,  or  ossicles,  are  named  the 
malleus,  incus,  and  stapes.  The  malleus  is  attached  by  a  long  slightly  curved 
process,  called  its  handle,  to  the  membrana  tympani,  the  line  of  attachment 


7O2  THE    SENSES 

being  vertical,  including  the  whole  length  of  the  handle,  and  extending  from 
the  upper  border  to  the  center  of  the  membrane.  The  head  of  the  malleus 
is  irregularly  rounded;  its  neck,  or  the  line  of  boundary  between  it  and  the 
handle,  supports  a  short  conical  process  which  receives  the  insertion  of  the 
tensor  tympani  muscle.  The  incus,  shaped  like  a  bicuspid  molar  tooth,  is 
articulated  by  its  broader  part  to  the  malleus.  Of  its  two  fang-like  processes,, 
one  directed  backward  has  a  free  end  lodged  in  a  depression  in  the  mastoid 
bone;  the  other,  curved  downward  and  more  pointed,  articulates  by  means 
of  a  roundish  tubercle  with  the  stapes.  The  stapes  is  a  little  bone  shaped 
exactly  like  a  stirrup,  of  which  the  base  or  bar  fits  into  the  f  enestra  ovalis. 
The  stapedius  muscle  is  attached  to  the  neck  of  the  stapes. 

The  bones  of  the  ear  are  covered  with  mucous  membrane  reflected  over 
them  from  the  wall  of  the  tympanum.  They  are  movable  both  altogether 
and  one  upon  the  other.  The  malleus  moves  and  vibrates  with  every  move- 


FIG.  433.  FIG.  434. 

FIG.  433. — Right  Bony  Labyrinth,  Viewed  from  the  Outer  Side.  The  specimen  here 
represented  is  prepared  by  separating  piecemeal  the  looser  substance  of  the  petrous  bone 
from  the  dense  walls  which  immediately  enclose  the  labyrinth,  i,  The  vestibule;  2, 
fenestra  ovalis;  3,  superior  semicircular  canal;  4,  horizontal  or  external  canal;  5,  posterior 
canal;  *,  ampullae  of  the  semicircular  canals;  6,  first  turn  of  the  cochlea;  7,  second  turn;  8, 
apex;  9,  fenestra  rotunda.  The  smaller  figure  in  outline  below  shows  the  natural  size. 
X  2.5.  (Sommering.) 

FIG.  434. — View  of  the  Interior  of  the  Left  Labyrinth.  The  bony  wall  of  the  labyrinth 
is  removed  superiorly  and  externally,  i,  Fovea  hemielliptica;  2,  fovea  hemispherica;  3, 
common  opening  of  the  superior  and  posterior  semicircular  canals;  4,  opening  of  the 
aqueduct  of  the  vestibule;  5,  the.superior,  6,  the  posterior,  and  17,  the  external  semicircular 
canals;  8,  spiral  tube  of  the  cochlea  (scala  tympani);  9,  opening  of  the  aqueduct  of  the 
cochlea;  10,  placed  on  the  lamina  spiralis  in  the  scala  vestibuli.  X  2 . 5.  (Sommering.) 

ment  and  vibration  of  the  membrana  tympani,  and  its  movements  are  com- 
municated through  the  incus  to  the  stapes,  and  through  the  stapes  to  the 
membrane  closing  the  fenestra  ovalis.  The  malleus,  also,  is  movable  in  its 
articulation  with  the  incus.  The  membrana  tympani  which  moves  the  long 
process  of  the  malleus  is  altered  in  its  degree  of  tension  by  the  degree  of  con- 


THE    INTERNAL    EAR 


705 


traction  of  the  tensor  tympani  muscles.  The  stapes  is  movable  on  the  proc- 
ess of  the  incus,  the  contractions  of  the  stapedius  muscle  draws  it  outward. 
The  axis  round  which  the  malleus  and  incus  rotate  is  the  line  joining  the 
processus  gracilis  of  the  malleus  and  the  posterior  process  of  the  incus. 

The  Internal  Ear. — The  internal  ear,  or  labyrinth,  constitutes  the 
proper  organ  of  hearing.  It  contains  special  epithelial  structures  to  which 
are  distributed  the  auditory  nerves.  The  organ  is  located  in  a  cavity  in  the 


FIG  435. — Membranous  Labyrinth  of  a  30  mm.  Human  Fetus.     A,  Viewed  from  its  Lateral 
Aspect;  B,  viewed  from  the  mesial  aspect.     (Streeter.) 


petrous  bone,  called  the  osseous  labyrinth.  The  auditory  organ  within  is 
called  the  membranous  labyrinth.  The  membranous  labyrinth  contains  a 
fluid  called  endolymph;  while  outside  it,  between  it  and  the  osseous  labyrinth, 
is  a  fluid  called  perilymph.  This  is  not  a  pure  lymph,  as  it  contains  mucin. 
The  osseous  labyrinth  consists  of  three  principal  parts,  namely  the  vesti- 
bule, the  cochlea,  and  the  semicircular  canals,  containing  the  respective 
divisions  of  the  membranous  labyrinth.  The  osseous  labyrinth  possesses 
openings  on  its  inner  wall  for  the  entrance  of  the  divisions  of  the  auditory 
nerve  from  the  cranial  cavity,  in  its  outer  wall  the  fenestra  ovalis,  2,  figure 
433,  an  opening  filled  by  the  base  of  the  stapes,  and  the  fenestra  rotunda. 
The  vestibule  also  presents  an  opening,  the  orifice  of  the  aqueductus  vestibuli* 


7°4  THE    SENSES 

The  Membranous  Labyrinth. — The  membranous  labyrinth  cor- 
responds generally  with  the  form  of  the  osseous  labyrinth,  so  far  as  regards 
the  vestibule  and  semicircular  canals,  but  is  separated  from  the  walls  of  these 
parts  by  perilymph,  except  where  the  nerves  enter  into  connection  within  it. 
The  labyrinth  is  a  closed  membrane  containing  endolymph. 

The  Utriculus  and  the  Sacculus. — The  vestibular  portion  of  the  inner  ear 
consists  of  membranous  sacs,  the  upper,  the  utriculus,  the  lower  called  the 
sacculus.  The  former  is  connected  with  the  semicircular  canals,  the  latter 
with  the  cochlea  by  the  cochlear  canal.  The  utriculus  and  the  sacculus  have 
on  their  floors  each  a  special  patch  of  sensory  epithelium  called  the  macula. 
The  fibers  of  the  vestibular  divisions  of  the  auditory  nerve  end  in  the 
maculae,  figure  435.  In  the  cavities  of  the  sacculus  and  utriculus  are  small 
masses  of  calcareous  particles  called  otoliths. 

The  Semicircular  Canals. — There  are  three  semicircular  canals  for  each 
ear,  one  horizontal  and  two  vertical  ones  placed  almost  at  right  angles  to 
each  other.  The  three  canals,  therefore,  occupy  the  three  planes  of  space. 
Each  has  a  considerable  enlargement  or  swelling,  called  an  ampulla.  The 
epithelium  of  the  ampulla  is  modified  at  the  point  of  entrance  of  the  nerve 
into  a  thickened  hillock  called  the  crista  acustica.  This  epithelium  is  com- 
posed of  rod  cells  or  supporting  cells  which  extend  the  full  thickness  of  the 

crista,   and  of  hair  cells,  which  occupy  the 
inner  or  free  half  of  the  crista.     The  hair 
cells  are  the  sensory  cells.     They  have  hair- 
like   processes  which  project  from  the  free 
ends  of  the  cells  out  into  the  endolymph  of 
the  cavity.     Nerve  fibrils  run  up  into  the 
FIG.  436.— View  of  the  Osseous     crista  and  apparently  form  terminal  arboriza- 
Cochlea    Divided    through    the     tions  at>out  the  hair  cells,  or,  according  to 
Middle,     i,  Central  canal  of  the 

modiolus;  2,  lamina  spiralis  ossea;     some  observers,  end  in  the  cells. 
3,  scala  tympani;  4,  scala  vestibuli;          The  Cochlea  and  the  Organ  of  Corti, — 

5,  porous  substance  of  the  modiolus  .       .     .  . 

near  one  of  the  sections  of  the  The  membranous  cochlea  is  located  in  the 
canalis  spiralis  modioli.  X  5.  spiral  canal  in  the  petrous  bone,  called  the 

cochlear  canal.      It  is  attached  to  the  wall 

of  the  cavity  between  the  fenestra  ovalis  and  the  fenestra  rotunda,  and  to 
the  outer  wall  of  the  canal  and  the  free  border  of  the  lamina  spiralis  almost, 
but  not  quite,  to  its  summit.  A  small  cavity  is  thus  left  around  the 
upper  end  of  the  cochlea  connecting  the  scala  vestibuli  above  with  the  scala 
tympani  below.  A  cross-section  through  the  cochlear  canal  shows  the 
relations  of  the  cochlear  canal  which  was  named  scala  media  by  the  earlier 
anatomists.  The  free  portion  of  the  membranous  wall  above  is  called  the 
membrane  of  Reisner,  while  that  below  is  called  the  basilar  membrane.  The 
basilar  membrane  supports  the  special  sensory  apparatus  for  the  reception 
of  stimuli  of  sound  waves. 


THE    COCHLEA   AND    THE   ORGAN   OF    CORTI 


705 


Organ  of  Corti. — The  basilar  membrane  supports  cells  of  several  types. 
About  midway  between  the  outer  edge  of  the  lamina  spiralis  and  the  outer 
wall  of  the  cochlea  are  situated  the  rods  of  Corti.  Viewed  sideways,  they  are 
seen  to  consist  of  an  external  and  internal  pillar,  each  rising  from  an  ex- 
panded foot  or  base  on  the  basilar  membrane,  figure  438.  They  slant  in- 
ward toward  each  other,  and  each  ends  in  a  swelling  termed  the  head,  the 
head  of  the  inner  pillar  overlying  that  of  the  outer,  figure  438.  Each  pair  of 
pillars  forms,  as  it  were,  a  pointed  roof  arching  over  a  space,  and  by  a  suc- 
cession of  them  a  little  tunnel  is  formed.  It  has  been  estimated  that  there 


FIG.  437. — Semidiagrammatic  Section  of  a  Cochlear  Whorl.     (After  Heitzmann.) 

are  about  four  thousand  of  these  pairs  of  rods  of  Corti  between  the  base  of 
the  cochlea  and  its  apex,  in  man  3,848  outer  and  5,590  inner  rods  (Retzius). 
They  are  found  progressively  to  increase  in  length,  and  become  more 
oblique;  in  other  words,  the  tunnel  becomes  wider,  but  diminishes  in  height 
as  we  approach  the  apex  of  the  cochlea. 

Leaning  against  the  rods  of  Corti  and  apparently  supported  by  them 
are  sensory  cells  or  hair  cells.  The  hair  cells  are  in  two  series,  the  inner  and 
the  outer  hair  cells.  The  former  consist  of  a  single  layer,  the  latter  of  three 
or  four  layers,  figure  438.  There  are  two  additional  types  of  supporting  cells, 
the  cells  of  Deiters  and  of  Hensen.  The  whole  structure  when  viewed  from 
above  bears  a  remarkable  resemblance  to  the  keyboard  of  a  piano. 

The  cochlear  division  of  the  auditory  nerve  enters  the  base  of  the  modiolus 
and  sends  a  spiral  whorl  of  fibers  out  under  the  spiral  lamina.  The  gan- 
glionic  cells  of  the  cochlear  division  of  the  auditory  nerve  are  located  in  the 

45 


706 


THE    SENSES 


base  of  the  lamina  where  they  form  the  spiral  ganglion.  The  nerve  fibers 
from  the  ganglion  cells  pass  out  through  small  holes  in  the  periphery  of  the 
spiral  plate  of  bone,  to  enter  the  organ  of  Corti.  Here  they  form  small 
longitudinal  bundles  that  quickly  end  about  the  hair  cells. 

THE  SENSE  OF  HEARING. 

All  the  acoustic  contrivances  of  the  organ  of  hearing  are  means  for  con- 
ducting sound.  Since  all  matter  is  capable  of  propagating  sonorous  vibra- 
tions, the  simplest  conditions  must  be  sufficient  for  mere  hearing;  since  all 
substances  surrounding  the  auditory  apparatus  would  stimulate  it.  The 
complex  development  of  the  organ  of  hearing,  therefore,  must  have  for  its 


mcmbrana  lectori* 


outer  hair-cells 


•    • 


inner  rod    vas    basilar         outer    cells  of  Deiters 
membrane     rod 


nerve  fbrc* 

FIG.  438. — Semidiagrammatic  Representation  of  the  Organ  of  Corti  and  Adjacent 
Structures.  (Merkel-Henle.)  a,  Cells  of  Hensen;  b,  cells  of  Claudius;  c,  internal  spiral 
sulcus;  x,  Nuel's  space.  The  nerve  fibers  (dendrites  of  cells  of  the  spinal  ganglion)  are 
seen  passing  to  Corti's  organ  through  openings  (foramina  nervosa)  in  the  bony  spiral 
lamina.  The  black  dots  represent  longitudinally  running  branches,  one  bundle  lying  to 
the  inner  side  of  the  inner  pillar,  a  second  just  to  the  outer  side  of  the  inner  pillar  within 
Corti's  tunnel,  the  third  beneath  the  outer  hair  cells. 

object  the  more  effective  propagation  of  the  sonorous  vibrations  and  their 
intensification  by  resonance;  and,  in  fact,  the  whole  of  the  acoustic  apparatus 
may  be  shown  to  have  reference  to  these  principles. 

The  external  ear  and  the  auditory  passages  influence  the  propagation  of 
sound  to  the  tympanum  by  collecting  from  the  atmosphere  the  sonorous 
undulations  that  strike  against  the  external  ear  and  by  transmitting  them  by 
the  air  in  the  passage  to  the  membrana  tympani. 

In  animals  living  in  the  atmosphere,  the  sonorous  vibrations  are  con- 
veyed to  the  auditory  epithelium  through  three  different  media  in  series; 
namely,  the  air  of  the  external  ear  and  meatus,  which  sets  in  vibration  the 
tympanic  membrane,  the  solid  chain  of  auditory  ossicles,  and  the  fluid  of  the 
labyrinth.  Sonorous  vibrations  are  imparted  too  imperfectly  from  air  to 


SOUND  707 

the  solid  structures  of  the  body  as  a  whole  for  the  propagation  of  sound  to 
the  internal  ear  to  be  adequately  effected  by  that  means  alone.  In  passing 
from  air  directly  into  water,  sonorous  vibrations  suffer  also  a  considerable 
diminution  of  their  strength;  but  if  a  tense  membrane  exists  between  the 
air  and  water,  the  sonorous  vibrations  are  communicated  from  the  former 
to  the  latter  medium  with  very  great  intensity.  This  fact,  of  which  Miiller 
gives  experimental  proof,  furnished  at  once  an  explanation  of  the  use  of  the 
fenestra  ovalis  and  of  the  membrane  closing  it.  It  is  the  means  of  com- 
municating, in  full  intensity,  the  vibrations  of  the  ear  bones,  or,  in  their 
absence,  of  the  air  in  the  tympanum,  to  the  fluid  of  the  labyrinth.  The 
vibration  of  the  fluids,  the  perilymph  and  endolymph,  of  the  internal  ear,  sets 
the  basilar  membrane  in  -vibration  and  in  consequence  stimulates  the  sensory 
apparatus  resting  upon  it.  This  last  is  the  essential  stimulating  act,  while 
all  that  precedes  is  more  or  less  accessory  or  contributory  to  this  act.  Just 
what  the  accessory  apparatus  contributes  can  be  best  understood  by  an  ex- 
amination of  the  stimulus  and  the  sensation  which  results  from  its  action. 

Sound. — Any  elastic  body,  e.g.,  air,  a  membrane,  or  a  string,  performing 
a  certain  number  of  regular  vibrations  per  second,  gives  rise  to  what  is  termed 
a  musical  sound  or  tone.  We  must,  however,  distinguish  between  a  musical 
sound  and  a  mere  noise;  the  latter  being  due  to  irregular  vibrations. 

Musical  sounds  are  distinguished  from  each  other  by  three  qualities: 
i.  Strength  or  intensity,  which  is  due  to  the  amplitude  or  length  of  the  wave 
of  vibrations.  2.  Rate,  the  number  of  vibrations  in  a  second.  3.  Quality, 
or  timbre,  the  peculiar  property  by  which  we  distinguish  the  same  note 
sounded  on  two  instruments,  e.g.,  a  piano  and  a  flute.  It  has  been  proved 
by  Helmholtz  to  depend  on  the  number  of  secondary  tones,  termed  harmonics, 
which  are  present  with  the  predominating  or  fundamental  tone;  that  is, 
rhythmic  vibrations  are  either  simple  in  form,  like  the  vibrations  of  a  reed 
or  tuning-fork,  or  compound,  like  the  vibrations  of  a  violin  or  piano  string. 
If  the  string  of  a  violin  is  plucked  it  not  only  vibrates  as  a  whole,  but  in  seg- 
ments in  the  ratio  of  one,  two,  three,  etc.  The  form  of  air  wave  that  is  pro- 
duced by  several  such  vibrating  bodies  is  very  complex  indeed,  as,  for  exam- 
ple, when  an  orchestra  is  playing. 

The  compound  wave  can  be  analyzed  into  its  constituent  elements  by  a 
system  of  resonators,  on  the  principle  of  sympathetic  vibration.  If  one 
sounds  a  series  of  musical  notes  before  such  a  system  of  resonators  it  will  be 
found  that  the  tones  and  overtones  are  selected  by  the  resonators  and  made 
more  prominent  so  that  they  can  be  identified. 

The  sensation  of  sound  has  in  it  certain  elements  that  correspond  closely 
with  the  physical  properties  of  sound,  i.e.,  loudness,  pitch,  and  quality. 
Loudness  is  dependent  merely  on  the  intensity  of  the  stimulation.  A  sound 
wave  of  great  energy,  for  example,  produces  a  larger  movement  of  the  tym- 
panic membrane,  and  it,  through  the  chain  of  bones  and  the  fluid  of  the 


708  THE   SENSES 

internal  ear,  a  larger  swing  of  the  basilar  membrane,  hence  a  more  intense 
stimulus  of  the  organ  of  Corti. 

Function  of  the  External  and  Middle  Ears. — It  has  already  been 
stated  that  the  external  ear  collects  the  sound  waves  and  conducts  them 
against  the  membrana  tympani.  This  membrane  vibrates  as  a  whole  to 
the  compound  waves  that  impinge  upon  it,  and  thus  serves  for  the  trans- 
mission of  sound  from  the  air  to  the  chain  of  ossicles  of  the  middle  ear.  It  is 
often  compared  to  the  membrane  of  a  drum,  but  there  are  fundamental 
differences. 

When  a  drum  is  struck,  a  certain  definite  fundamental  tone  is  elicited; 
similarly  a  drum  is  thrown  into  vibration  when  certain  tones  are  sounded  in 
its  neighborhood,  while  it  is  quite  unaffected  by  others.  In  other  words,  it 
can  take  up  and  vibrate  in  response  to  only  those  tones  whose  vibrations 


B 


FIG.  439. — Showing  A  and  B,  Simple  Pendular  Vibrations,  Separated  by  One  Octave.     C 
The  form  of  the  curve  produced  by  the  combination  of  A  and  B. 

correspond  in  number  with  those  of  its  own  fundamental  tone.  The 
tympanic  membrane  can  vibrate  in  response  to  a  wide  range  of  tones,  thus 
transmitting  vibration  frequencies  ranging  between  30  to  4,000  or  5,000 
per  second,  or  even  higher.  The  limits  of  hearing  as  regards  pitch  are  in 
the  ordinary  average  ear  represented  by  tones  of  these  vibration  fre- 
quencies. In  some  individuals  the  scale  extends  to  30,000  or  more.  This 
range  is  made  possible  by  the  fact  that  the  membrana  tympani  is  by  no 
means  under  constant  tension,  but  is  connected  with  the  chain  of  auditory 
ossicles,  the  contractions  of  the  muscle  of  which  produce  variations  in  ten- 
sion, especially  on  the  handle  of  the  malleus.  If  the  membrane  were  quite 
free  in  its  center,  it  would  be  like  a  drum  which  goes  on  vibrating  for  some 
time  after  it  is  struck,  and  each  sound  would  be  prolonged,  leading  to  con- 
siderable confusion.  This  evil  is  obviated  by  the  ear  bones,  which  check 
the  continuance  of  the  vibrations  like  the  "dampers"  in  a  piano. 

The  vibrations  of  the  membrana  tympani  are  transmitted  by  the  chain 
of  ossicles  to  the  fenestra  ovalis  and  the  fluid  of  the  labyrinth,  their 


FUNCTION    OF   THE   EXTERNAL   AND    MIDDLE    EARS 


709 


dispersion  in  the  tympanum  being  prevented  by  the  difficulty  of  the 
transition  of  vibrations  from  solid  to  gaseous  bodies.  The  necessity  of  the 
presence  of  air  on  the  inner  side  of  the  membrana  tympani  is  obvious. 
Without  this  provision,  neither  would  the  vibrations  of  the  membrane  be 
free  nor  the  chain  of  bones  isolated  so  as  to  propagate  the  sonorous  undula- 
tions with  concentration  of  their  intensity.  While  the  oscillations  of  the 
membrana  tympani  are  readily  communicated  to  the  air  in  the  cavity  of 
the  tympanum,  those  of  the  solid  ossicles  will  not  be  conducted  away  by 
the  air,  but  will  be  propagated  to  the  labyrinth  without  being  dispersed  in 
the  tympanum.  The  existence  of  the  membrane 
over  the  fenestra  rotunda  permits  vibration  of 
the  stapes  to  and  from  the  labyrinth.  When 
the  membrane  of  the  fenestra  ovalis  is  pressed 
toward  the  labyrinth  by  the  stapes,  the  pressure 
is  communicated  through  the  fluid  of  the  laby- 
rinth toward  the  cavity  of  the  tympanum  at 
the  fenestra  rotunda  which  yields. 

The  propagation  of  sound  through  the 
auditory  ossicles  to  the  labyrinth  must  be 
effected  by  oscillations  of  the  bones  as  a  whole. 
The  long  process  of  the  malleus  receives  the 
undulations  of  the  membrana  tympani,  figure 
440,  a,  a,  in  a  direction  indicated  by  the  arrows 
and  nearly  perpendicular  to  itself.  The  vibra- 
tions of  the  long  process  of  the  malleus  sets  the 
chain  of  bones  vibrating  about  the  axis  fixed  by 
the  attachment  of  the  anterior  ligament  of  the 
malleus  and  the  ligaments  of  the  incus,  see 
figure  432.  From  the  long  process  of  the  incus,  which  is  parallel  with  the 
long  process  of  the  malleus,  the  undulations  are  communicated  to  the 
stapes  and  by  the  stapes  to  the  fenestra  ovalis  in  a  perpendicular  direc- 
tion. Increasing  tension  of  the  membrana  tympani  diminishes  the  facility 
of  transmission  of  sonorous  undulations  from  the  air  to  it.  It  has  been 
inferred,  therefore,  that  hearing  is  rendered  less  acute  by  increasing  the 
tension  of  the  membrana  tympani.  This  is  accomplished  by  the  contrac- 
tions of  the  tensor  tympani  muscle.  The  exact  influence  of  the  stapedius 
muscle  in  the  act  of  hearing  is  unknown.  It  acts  upon  the  stapes  in  such 
a  manner  as  to  make  it  rest  obliquely  in  the  fenestra  ovalis,  depressing 
that  side  of  the  stapes  on  which  it  is  attached  and  elevating  the  other 
side  to  the  same  extent.  It  seems  to  prevent  too  great  a  movement  of 
the  bone. 

The  pharyngeal  orifice  of  the  Eustachian  tube  is  usually  shut.     During 
swallowing,  however,  it  is  opened;  which  may  be  shown  as  follows:     If  the 


FIG.  440. — Diagram  to  Il- 
lustrate the  Action  of  the 
Ossicles  of  the  Middle  Ear  in 
the  Conduction  of  Sound  to 
the  Internal  Ear. 


710  THE    SENSES 

nose  and  mouth  be  closed  and  the  cheeks  blown  out,  a  sense  of  pressure  is 
produced  in  both  ears  the  moment  we  swallow.  This  is  due,  doubtless,  to 
the  bulging  out  of  the  tympanic  membrane  by  the  compressed  air,  which  at 
that  moment  enters  the  Eustachian  tube.  The  principal  office  of  the  Eusta- 
chian  tube  has  relation  to  the  prevention  of  the  effects  of  increased  tension  of 
the  membrana  tympani.  Its  existence  and  openness  will  provide  for  the 
maintenance  of  the  equilibrium  between  the  air  within  the  tympanum  and 
the  external  air,  so  as  to  prevent  the  inordinate  tension  of  the  membrana 
tympani  which  would  be  produced  by  too  great  or  too  little  pressure  on  either 
side.  While  discharging  this  office  it  serves  as  an  outlet  for  mucus.  If  the 
tube  were  permanently  open,  the  sound  of  one's  own  voice  would  probably 
be  greatly  intensified,  a  condition  which  would  of  course  interfere  with  the 
perception  of  other  sounds.  At  any  rate,  it  is  certain  that  sonorous  vibra- 
tions can  be  propagated  up  the  tube  to  the  tympanum  by  means  of  a  catheter 
inserted  into  the  pharyngeal  orifice  of  the  Eustachian  tube. 

The  Function  of  the  Internal  Ear. — The  fluids  of  the  labyrinth  re- 
ceive the  sonorous  vibrations  at  the  fenestra  ovalis  and,  we  must  assume, 
conduct  the  same  throughout  the  cavity.  In  all  forms  of  organs  of  hearing 
even  to  the  simplest,  liquid  is  the  medium  through  which  the  auditory  sensory 
epithelium  is  stimulated.  We  have  already  seen  that  in  the  mammalian  ear 
there  is  a  special  mechanical  arrangement  to  intensify  the  vibrations  of  the 
fluid  in  the  cochlear  canal. 

The  utriculus,  sacculus,  and  semicircular  canals  are  probably  not  con- 
cerned with  auditory  function,  but  with  the  sense  of  equilibrium;  hence  they 
will  be  discussed  separately  a  little  later. 

The  cochlea  is  the  special  organ  of  hearing.  When  it  is  set  in  vibration 
the  movement  stimulates  the  sensory  hair  cells  on  the  basement  membrane, 
producing  a  sensory  impulse  which  is  transmitted  along  the  paths  to 
the  brain  and  there  produces  an  auditory  sensation.  If  the  stimulus 
results  from  a  disturbance  of  an  explosive  or  non-harmonic  nature,  the 
sensation  is  interpreted  as  a  noise.  If  the  disturbance  is  rhythmic  or 
harmonic  and  repeated  in  sequence  within  certain  limits  of  rate,  then  a 
tone  is  perceived. 

The  intensity  of  sound,  the  energy  of  the  disturbance,  affects  the  basilar 
membrane  by  producing  motion  of  varying  amplitude.  This  stimulates  the 
hair  cells  with  greater  or  less  intensity,  which  can  be  detected  by  the  sensorium 
as  loudness.  Loudness  of  the  sound  sensation  is  interpreted  as  intensity  of 
sound  wave. 

The  interpretation  of  pitch  is  accomplished  by  the  ear  through  a  wide 
range  of  rates  of  vibration  that  produce  sensations  of  tone.  The  average 
person  can  perceive  musical  tones  over  a  range  of  vibration  of  from  sixty-four 
double  vibrations  per  second  for  the  lower  notes,  to  four  thousand  and  ninety- 
six  for  the  higher  notes.  These  limits  may  be  extended  to  thirty  per  second 


THE   FUNCTION   OF    THE   INTERNAL   EAR  71 1 

and  forty  thousand  per  second,  respectively,  but  only  a  small  number  of 
tones  can  be  perceived  outside  of  the  narrower  limits  given  above.  This 
extraordinary  range  of  tone  is  conceivable  only  on  the  supposition  of  local- 
ization of  the  stimulus  in  some  part  of  the  organ.  Most  physiologists  look 
to  the  basilar  membrane  and  the  organ  of  Corti  for  the  localization. 

Suppose  a  simple  tuning-fork  to  be  vibrating  with  a  frequency  of  sixty- 
four  per  second,  then  these  waves  will  be  conducted  through  the  auditory 
apparatus  until  they  fall  on  the  basilar  membrane,  and  will  set  it  in  vibration 
at  the  same  rate.  The  exact  type  of  the  vibration  is  at  present  a  matter  of 
inference.  The  piano  theory  of  Helmholtz  gives  probably  the  most  satisfac- 
tory explanation.  It  assumes  that  the  basilar  membrane  vibrates  as  would 
a  number  of  strings  set  in  its  transverse  dimension.  In  support  of  this  as- 
sumption it  is  asserted  that  the  membrane  is  taut  in  the  transverse  and 
loose  in  the  longitudinal  plane.  Retzius  has  estimated  that  it  contains 
about  24,000  fibers,  and  that  it  measures  in  width  at  the  base  0.135  mm- 
and  at  the  apex  o .  234  mm.  In  the  above  illustration  the  vibration  fre- 
quency of  sixty-four  would  supposedly  set  in  sympathetic  vibration  that  part 
of  the  apex  of  the  basilar  membrane  which  vibrated  in  the  same  frequency, 
and  the  sensory  cells  of  the  organ  of  Corti,  located  over  the  vibrating  fiber, 
would  be  stimulated  accordingly.  In  the  same  way  notes  of  medium  and 
of  high  frequency  stimulate  localized  areas  of  sensory  cells  in  the  middle 
and  basal  parts  of  the  organ  of  Corti  and  produce  sensations  of  correspond- 
ing pitch. 

This  idea  of  localization  of  auditory  sensory  stimulation  makes  it  easier 
to  understand  the  analysis  by  the  ear  of  compound  sonorous  waves.  Such 
waves  impinge  on  the  membrana  tympani  and  are  transmitted  through  the 
conducting  media  unanalyzed,  and  may  be  supposed  to  fall  on  the  basilar 
membrane  as  compound  waves.  The  basilar  fibers  acting  like  so  many 
resonators,  take  up  the  constituent  sonorous  elements  in  sympathetic  vibra- 
tion. In  short,  the  basilar  membrane  is  an  analyzer  in  which  the  compound 
wave  is  reduced  to  its  simple  components,  each  of  which  stimulates  its  cor- 
responding portion  of  the  organ  of  Corti.  The  auditory  nerve  impulses  are 
conducted  through  the  cochlear  nerves  to  the  sensorium  where  they  produce 
auditory  sensations  with  the  same  definiteness  of  pattern  as  cutaneous  or 
optical  stimuli  produce  sensations  that  correspond  to  the  patterns  of  stimula- 
tion. The  audition  is  so  definite  that  one  can  consciously  pick  out  one  or 
the  other  of  the  constituent  stimulating  elements  and  follow  and  examine 
the  same  to  the  exclusion  of  the  others,  as  when  one  follows  a  single  instru- 
ment in  an  orchestra  or  a  single  voice  in  a  group  of  chattering  children. 

Bernstein  says  of  this  wonderful  organ: 

"In  the  cochlea  we  have  to  do  with  a  series  of  apparatus  adapted  for  per- 
forming sympathetic  vibrations  with  wonderful  exactness.  We  have  here 
before  us  a  musical  instrument  which  is  designed  not  to  create  musical 


7*2  THE    SENSES 

sounds,  but  to  render  them  perceptible,  and  which  is  similar  in  construction 
to  artificial  musical  instruments,  but  which  far  surpasses  them  in  the  delicacy 
as  well  as  the  simplicity  of  its  execution.  For,  while  in  a  piano  every  string 
must  have  a  separate  hammer  by  means  of  which  it  is  sounded,  the  ear  pos- 
sesses a  single  hammer  of  an  ingenious  form  in  its  ear  bones,  which  can  make 
every  string  of  the  organ  of  Corti  sound  separately." 

Auditory  Judgments.  —  Direction. — The  power  of  perceiving  the 
direction  of  sounds  is  not  a  faculty  of  the  sense  of  hearing  itself,  but  is  an  act 
of  the  mind  judging  by  experience  previously  acquired.  From  the  modifica- 
tions which  the  sensation  of  sound  undergoes  according  to  the  direction  in 
which  the  sound  reaches  us,  the  mind  infers  the  position  of  the  sounding 
body.  The  only  true  guide  for  this  inference  is  the  more  intense  action  of  the 
sound  upon  one  than  upon  the  other  ear.  But  even  here  there  is  room  for 
much  deception,  by  the  influence  of  reflexion  or  resonance,  and  by  the  prop- 
agation of  sound  from  a  distance,  without  loss  of  intensity,  through  curved 
conducting  tubes  filled  with  air.  By  means  of  such  tubes,  or  of  solid  con- 
ductors, which  convey  the  sonorous  vibrations  from  their  source  to  a  distant 
resonant  body,  sounds  may  be  made  to  appear  to  orginate  in  a  new  situation. 
The  direction  of  sound  may  also  be  judged  of  by  means  of  one  ear  only;  the 
position  of  the  ear  and  head  being  varied,  so  that  the  sonorous  undulations 
at  one  moment  fall  upon  the  ear  in  a  perpendicular  direction,  at  another 
moment  obliquely.  But  when  neither  of  these  circumstances  can  guide  us  in 
distinguishing  the  direction  of  sound,  as  when  it  falls  equally  upon  both  ears, 
its  source  being,  for  example,  either  directly  in  front  or  behind  us,  it  becomes 
impossible  to  determine  whence  the  sound  comes. 

Distance. — The  judgment  of  the  distance  of  the  source  of  sounds  is  in- 
ferred from  their  intensity.  The  sound  is  interpreted  as  coming  from  an 
exterior  sonorous  body.  When  the  intensity  of  the  voice  is  modified  in  imita- 
tion of  the  effect  of  distance,  it  excites  the  idea  of  its  originating  at  a  distance. 
Ventriloquists  take  advantage  of  the  difficulty  with  which  the  direction  of 
sound  is  recognized,  and  also  the  influence  of  the  imagination  over  our  judg- 
ment, when  they  modulate  the  voices,  and  at  the  same  time  pretend,  them- 
selves, to  hear  sounds  as  coming  from  a  certain  direction. 

Duration  of  the  Auditory  Stimulus. — By  removing  one  or  several  teeth 
from  the  toothed  wheel  of  a  vibrator,  the  fact  has  been  demonstrated  that  in 
the  case  of  the  auditory  organ,  as  in  that  of  the  eye,  the  sensation  continues 
longer  than  the  impression  which  causes  it;  for  a  removal  of  the  tooth  pro- 
duced no  interruption  of  the  sound.  The  gradual  cessation  of  the  sensation 
of  sound  renders  it  difficult  to  determine  its  exact  duration  beyond  that  of 
the  impression  of  the  sonorous  impulses. 

Binaural  Sensations. — Sound  stimulates  each  of  the  two  ears  yet  ordi- 
narily only  one  sound  sensation  is  perceived,  just  as  in  binocular  vision 
,only  a  single  object  is  seen.  Analogous  to  the  double  vision  dependent  on 


THE    SENSE    OF   EQUILIBRIUM  713 

unequal  refraction  is  the  double  hearing  of  a  single  sound  owing  to  the 
sound  coming  to  the  ear  through  media  of  unequal  conducting  power. 
The  phenomenon  depends  on  the  unequal  conducting  power  of  two  media 
through  which  one  and  the  same  sound  is  transmitted  to  the  ear.  If  a 
small  bell  be  sounded  in  water  while  the  ears  are  closed  by  plugs,  and  a 
solid  conductor  be  interposed  between  the  water  and  one  ear,  two  sounds 
will  be  heard  differing  in  intensity  and  tone,  one  being  conveyed  to  the 
ear  through  the  medium  of  the  atmosphere,  the  other  through  the  con- 
ducting-rod. 

Subjective  Sensations. — Subjective  sounds  are  the  result  of  a  state  of  irri- 
tation or  excitement  of  the  auditory  nerve  produced  by  other  causes  than 
sonorous  impulses.  A  state  of  excitement  of  this  nerve  or  its  tracts 
gives  rise  to  the  sensation  of  sound,  hence  the  ringing  and  buzzing  in  the 
ears  heard  by  persons  of  irritable  and  exhausted  nervous  system,  and  by 
patients  with  cerebral  disease,  or  disease  of  the  auditory  nerve  itself;  hence 
also  the  noise  in  the  ears  heard  for  some  time  after  a  long  journey  in  a  rat- 
tling, noisy  vehicle.  Ritter  found  that  electric  currents  also  excite  sounds  in 
the  ears.  From  the  above  truly  subjective  sound  we  must  distinguish  those 
dependent,  not  on  a  state  of  the  auditory  nerve  itself  merely,  but  on  sonorous 
vibrations  excited  in  the  auditory  apparatus.  Such  are  the  buzzing  sounds 
attendant  on  vascular  congestion  of  the  head  and  ear  or  on  aneurysmal  dilata- 
tion of  the  vessels.  Frequently  even  the  simple  pulsatory  circulation  of  the 
blood  in  the  ear  is  heard.  To  the  sounds  of  this  class  belong  also  the  buzz 
or  hum  heard  during  the  contraction  of  the  palatine  muscles  in  the  act  of 
yawning,  during  the  forcing  of  air  into  the  tympanum  so  as  to  make  tense  the 
membrana  tympani. 

Irritation  or  excitement  of  the  auditory  nerve  may  give  rise  to  move- 
ments in  the  body  and  to  sensations  seemingly  from  other  organs  of  sense. 
In  both  cases  it  is  probable  that  associated  centers  of  the  brain  stem  and 
connected  areas  of  the  cortex  come  into  play.  An  intense  and  sudden  noise 
excites  in  every  person  closure  of  the  eyelids,  and  in  nervous  individuals 
a  start  of  the  whole  body  or  an  unpleasant  sensation  through  the  body 
like  that  produced  by  an  electric  shock. 


THE  SENSE  OF  EQUILIBRIUM. 

Although  the  utriculus,  sacculus,  and  semicircular  canals  form  the  major 
part  of  the  labyrinth  and  are  closely  associated  with  the  cochlea  in  develop- 
ment, there  is  increasing  evidence  that  these  structures  are  not  concerned 
with  hearing,  but  rather  with  a  sense  of  equilibrium.  This  view  has  been 
strengthened  by  recent  investigations  into  the  anatomical  relations  of  the 
different  elements  in  the  auditory  nerve,  figure  435. 


7X4  THE    SENSES 

These  structures  have  each  a  special  modification  of  the  sensory  epithe- 
lium which  receives  the  vestibular  branch  of  the  eighth  nerve.  These 
epithelial  areas  are  differentiations  of  the  embryonic  ear  pit,  which  is  derived 
from  the  epiblast.  In  fishes  which  have  well-developed  semicircular  canals 
and  vestibule  this  sensory  epithelium  has  a  common  origin  from  the  em- 
bryonic anlage  which  gives  rise  to  the  ear,  the  branchial  sense  organ,  and 
the  lateral  line  organs,  all  of  which  probably  have  static  functions. 

The  Semicircular  Canals. — The  semicircular  canals  are  connected 
with  the  utriculus,  are  three  in  number  on  each  side,  and  have  been  already 
shown  to  lie  in  space  practically  at  right  angles  to  one  another.  Each  is 
filled  with  endolymph,  and  each  has  a  special  organ,  the  crista  acustica, 
which  receives  a  division  of  the  vestibular  branch  of  the  eighth  nerve. 

The  function  of  the 'semicircular  canals  has  long  been  believed  to  be  to 
give  rise  to  sensations  by  which  we  determine  the  motions  of  the  body  in 
space.  But  Maxwell  has  shown  that  motion  and  position  are  both  medi- 
ated by  the  combined  reactions  of  the  labyrinthine  sense  organs.  It  was 
shown  long  ago  that  if  one  closes  his  eyes  and  turns  rapidly  around  the  ver- 
tical axis,  then  suddenly  stops  and  opens  the  eyes,  surrounding  objects 
seem  to  be  rotating  around  the  same  vertical  axis.  If  the  face  is  in  the 
horizontal  plane  and  the  rotation  around  the  vertical  axis,  then,  upon 
suddenly  raising  the  head  into  the  ordinary  position  and  opening  the  eyes, 
objects  seem  to  be  moving  about  the  head  around  the  horizontal  axis.  In 
both  these  cases  the  direction  of  the  apparent  motion  of  objects  depends 
upon  the  actual  motion  of  the  body  that  preceded  it  and  is  in  the  opposite 
direction.  In  the  first  case  the  rotation  is  in  the  plane  of  the  horizontal 
semicircular  canal.  It  is  assumed  here  that,  at  the  beginning  of  such  a 
movement,  the  endolymph,  being  fluid  and  inert,  tends  to  remain  still  for  a 
moment  and  the  effect  is  to  produce  an  increase  in  pressure  in  the  funnel 
of  the  ampulla.  This  relative  increase  of  tension  on  the  hair  cells  of  the 
crista  acustica,  stimulates  the  hair  cells  and  gives  rise  to  sensory  nerve 
impulses.  When  the  head  suddenly  stops  rotating  the  situation  is  just 
reversed  and  stimulation  is  in  the  opposite  direction.  Considering  the 
position  of  the  three  semicircular  canals,  it  will  be  seen  that  movement  of 
the  head  in  any  direction  will  stimulate  one  or  more  of  the  cristae,  giving 
rise  to  either  simple  or  complex  sensory  impulses.  Stimuli  on  the  recessus 
utriculi  adds  to  the  effect. 

This  theory  is  borne  out  by  the  effects  of  operation  and  stimulation  of 
the  exposed  semicircular  canals.  Injury  to  the  semicircular  canals  causes 
disturbances  in  muscular  co-ordination,  especially  in  movements  that  take 
place  in  the  plane  of  the  injured  canal.  If  a  horizontal  canal  in  a  pigeon 
be  sectioned,  the  animal  supports  its  head  in  the  vertical  position  very 
well,  but  is  unable  to  co-ordinate  its  horizontal  movements.  It  tends  to 
produce  rotary  motions  around  the  vertical  axis.  If  a  vertical  canal  is  sec- 


THE    EYE  715 

tioned,  the  head  falls  to  one  or  the  other  side  according  to  the  canal,  and  the 
animal  shows  instability  of  position  in  that  plane.  It  has  been  shown  that 
stimulation  of  a  sectioned  canal  produced  reflex  movements  in  that  plane. 

Muscular  co-ordination  is  a  complex  phenomenon  and  involves  the  oper- 
ation of  numerous  sensory  impulses  from  other  organs  of  the  body,  especially 
from  the  eyes  and  general  skin.  Some  confusion  has  arisen  from  the  fact  that 
there  are  associated  with  the  disturbance  in  the  semicircular  canals  move- 
ments of  the  eyes  and  head  in  higher  animals,  and  of  the  eyes,  head,  and  fins 
in  such  animals  as  fishes — the  so-called  compensatory  movements.  Without 
going  into  details  it  is  sufficient  to  state  that  the  sense  organs  of  the  semi- 
circular canals  probably  form  only  one  of  the  series  of  sensory  structures 
concerned  in  the  co-ordination  of  body  movements. 

The  Utriculus  and  Sacculus. — The  utriculus  and  sacculus  each  have 
a  sensory  area,  the  maculae,  over  which  there  rests  in  the  human  ear 
and  in  most  animals  small  particles  of  calcareous  matter,  otoliths.  These 
otoliths,  therefore,  lie  among  the  projecting  hairs  of  the  sensory  cells.  This 
is  characteristic  of  these  sensory  areas  and  differentiates  them  from  the 
arrangement  present  in  the  cristae.  There  would  seem  to  be  close  agree- 
ment in  function  between  the  maculae  and  cristae,  and  we  naturally  look  to 
the  influence  of  the  otoliths  on  the  processes  which  result  in  the  stimulation 
of  the  maculae.  Attempts  have  been  made  to  remove  the  otoliths,  with  the 
result  that  in  such  animals  there  is  apparent  inability  to  maintain  a  constant 
position  in  space.  The  experiments  have  been  performed  which  have  sug- 
gested the  theory  that  the  otoliths  take  an  active  part  in  stimulating  the  sen- 
sory cells,  probably  by  their  mere  pressure.  If  the  head  is  inclined  in  one 
or  the  other  direction,  the  pressure  of  the  otoliths  will  shift  on  the  hair  cells, 
and  that  is  sufficient  to  stimulate  them.  If  this  view  is  correct,  then  we  may 
regard  these  structures  as  static  in  function  as  compared  with  the  semicircular 
canals,  which  are  dynamic.  The  anatomical  separation  of  the  nerves  for  the 
cochlea  from  the  division  for  the  utriculus,  sacculus,  and  semicircular  canals 
itself  suggests  isolation  in  function,  figures  389  and  435.  It  is  conceivable 
that  loud  noises  of  an  explosive  nature  may  cause  sufficient  vibration  of  the 
endolymph  to  affect  the  otoliths  and  thus  stimulate  the  cristae.  Yet,  if  such 
stimulation  takes  place  it  is  probably  only  of  secondary  importance. 


THE  SENSE  OF  SIGHT. 
THE  EYE. 

The  eye,  the  organ  of  vision,  is  the  most  complex  and  most  highly  devel- 
oped of  the  organs  of  special  sense.  It  consists  not  only  of  a  special  sen- 
sory epithelium,  the  retina,  sensitive  to  light  stimulation,  but  of  a  series  of 
special  structures  which  intensify  and  localize  the  stimulus.  There  are 
also  accessory  organs  for  the  protection  of  the  eye. 


7l6  THE    SENSES 

The  Eyelids  and  Lachrymal  Apparatus. — The  eyeball  is  kept  moist 
over  its  free  surface  and  protected  from  external  injury  by  the  eyelids, 
by  the  glands  that  secrete  the  lachrymal  fluid  to  moisten  the  surface  of  the 
cornea,  and  by  the  oil  glands  that  secrete  oil  on  the  margins  of  the  lids. 

The  conjunctiva,  or  lining  membrane  of  the  lids,  which  is  reflected  on  to 
the  free  surface  of  the  eyeball,  protects  the  eye  from  injury  by  its  extraor- 
dinary sensitiveness  to  irritation  by  dust  or  other  external  substance.  Its 
stimulation  produces  reflex  secretion  of  the  lachrymal  fluid  that  flows  over 
the  surface  of  the  eye,  and  tends  to  wash  away  the  stimulating  substance. 


FIG.  441. — Section  of  the  Eyeball. 

The  Eyeball  and  its  Parts. — The  detail  of  the  structure  of  the  eye- 
ball is  too  complex  to  be  given  here  except  in  so  far  as  seems  necessary  for  a 
clearer  presentation  of  the  physiological  facts.  A  gross  dissection  reveals 
the  tough,  white,  outer  coat,  the  sdera;  the  intermediate  thin,  vascular 
pigmented  coat,  the  choroid;  and  the  inner  nervous  coat,  the  retina. 

The  section  also  shows  that  the  eyeball  is  specialized  in  structure  in  its 
anterior  region  and  that  its  contained  cavity  is  divided  into  two  parts,  viz., 
the  anterior  and  posterior  chambers.  The  anterior  chamber  is  filled  with  the 
transparent  aqueous  fluid.  This  fluid  is  like  lymph  in  its  composition. 
The  posterior  chamber  between  the  lens  and  the  retina  is  filled  with  the 
clear  jelly-like  vitreous  substance. 

The  Cornea. — The  sclera  is  continuous  with  the  cornea  in  front  of  the 
eyeball,  but  the  cornea  is  transparent  and  its  radius  of  curvature  is  less  than 
that  of  the  main  portion  of  the  eye.  It  is  composed  of  stratified  epithelial 
cells,  and  is  richly  supplied  with  sensory  nerves  that  form  an  intraepithelial 


THE    CILIARY    APPARATUS    AND    THE    IRIS 


717 


plexus  of  delicate  varicose   f  brils.     The  cornea  has  no  blood  vessels,  but 
has  a  rich  network  of  lym-  .iatic  spaces. 

The  Lens. — The  ler  3  is  a  special  modification  of  epithelium  composed 
of  highly  refractive  material,  situated  just  behind  the  iris.  It  is  enclosed 
in  a  capsule  and  supported  in  its  place  by  the  suspensory  ligament,  which 


FIG.  442. — Vertical  Section  of  Rabbit's  Cornea,  a,  Epithelium  of  cornea,  showing  the 
different  shapes  of  the  cells  at  various  depths  from  the  free  surface;  &,  portion  of  the  proper 
substance  of  the  cornea.  (Klein.) 

is  fused  into  the  capsule  around  its  equator.  The  lens  is  a  biconvex  struc- 
ture composed  of  transparent  fibers  which  are  arranged  in  concentric 
layers.  Its  posterior  curvature  is  greater  than  the  anterior,  the  radii  being 
6  and  10  mm.  respectively. 


FIG.  443. 


FIG.  444. 


FIG.  443. — Ciliary  Processes,  as  Seen  from  Behind,  i,  Posterior  surface  of  the  iris, 
with  the  sphincter  muscle  of  the  pupil;  2,  anterior  part  of  the  choroid  coat;  3,  one  of  the 
ciliary  processes,  of  which  about  seventy  are  represented.  X  $. 

FIG.  444. — Laminated  Structure  of  the  Crystalline  Lens.  The  laminae  are  split  up 
after  hardening  in  alcohol,  i  The  denser  central  part  or  nucleus;  2,  the  successive  exter- 
nal layers.  X  4.  (Arnold.) 


The  Ciliary  Apparatus  and  the  Iris. — These  structures  are  a  con- 
tinuation and  modification  of  the  choroid  coat  in  the  anterior  portion  of  the 
eye.  Around  the  circumference  of  the  cornea  the  choroid  coat  is  consider- 
ably thickened  and  folded  in  the  modification  known  as  the  ciliary  apparatus. 


7i8 


THE    SENSES 


A  radial  layer  of  muscle,  figure  445,  is  knitted  into  the  base  of  the  cornea,  on 
the  one  hand,  and  extends  back  into  the  choroid,  on  the  other.  Thick  bun- 
dles of  the  circular  fibers  are  also  present  in  this  mass  of  muscle.  From  the 
ciliary  processes  extending  over  the  lens  is  the  iris.  It  is  a  sheet  of  connec- 
tive tissue  and  muscle  lined  with  epithelium  and  highly  pigmented. 

In  the  middle  anterior  portion  of  the  iris  is  a  round  aperture,  the  pupil. 
The  muscle  fibers  are  of  the  unstriated  muscle  type  and  are  arranged  in 
two  layers  one  circularly  and  the  other  radially.  Contractions  of  the  cir- 
cular muscles  of  the  iris  produce  constriction  of  the  pupil,  while  contrac- 
tions of  the  radial  fibers  produce  dilatation.  Both  the  ciliary  apparatus 
and  the  iris  are  supplied  with  motor  nerves. 


anterior  ciliary  arteries  and  veins 

greater  arterial  circle 
angle  of  the  Iris 

canal  of  schlemm 


ciliary  muscle 


limbus  corneae 


anterior  chamber 


epithelium 


anterior   limiting! 
membrane         f 


posterior  chamber 
epithelium  of  lens 


capsule  of  lens 


posterior  limiting  membrane 


stroraa  of  Iris 
posterior  surface  of  lite 
sphincter  of  pupil 


FIG.  445. — Meridional  Section  of  a  Portion  of  the  Anterior  Part  of  the  Eyeball.     (Toldt.) 

Fibers  of  the  third  cranial  nerve  are  distributed  to  the  ciliary  muscles, 
apparently  to  both  radial  and  circular  muscles,  and  when  these  nerves  are 
stimulating  the  resulting  contractions  of  the  muscles  tend  to  remove  the  ten- 
sion from  the  capsule  of  the  lens.  These  nerve  fibers  pass  through  the  ciliary 
ganglion  where  they  form  synapses  with  the  ganglionic  cells.  Motor  fibers 
from  the  third  cranial  nerve  also  supply  the  circular  muscle  of  the  iris,  which 
produces  constriction  of  the  pupil.  The  radial  muscles  of  the  iris  are 


STRUCTURE    OF    THE    RETINA 


719 


supplied  by  nerves  by  way  of  the  cervical  sympathetic,  superior  cervical 
ganglion,  and  the  ophthalmic  branch  of  the  fifth  cranial  nerve. 

Structure  of  the  Retina. — The  retina  occupies  the  deeper  half  of  the 
cup  of  the  eyeball.  It  extends  forward  as  far  as  the  ora  serrata,  where  its 
complex  structure  changes  the  form  to  a  simple  epithelial  layer,  which  lines 
the  anterior  portion  of  the  eyeball  and  the  ciliary  processes.  In  the  center 
of  the  retina  is  a  round  yellowish  spot  having  a  minute  depression  in  its 
center,  called  the  yellow  spot  of  Sommering.  The  depression  in  its  center  is 


Layer  of 

rods  and 

cones 

Mem- 

brana 

limitans 

externa 


Outer 

molecular 

layer 


]  Inner 
\  molecular 
layer 

Gangli- 

onic 

layer 

I  Stratum 
opticurfi 

Membrana  limitans  interna 
FIG.  446. — Section  of  Human  Retina.     (Cunningham,  modified  from  Schulze.) 

thefovea  centralis.  About  2 . 5  mm.  to  the  inner* side  of  the  yellow  spot  is  the 
point  at  which  the  optic  nerve  enters  and  spreads  out  its  fibers  into  the  retina. 
The  optic  nerve  arises  from  the  base  of  the  brain  and  passes  forward 
toward  the  orbit,  being  covered  by  the  membranes  which  cover  the  brain. 
The  fibers  of  the  optic  nerve  are  exceedingly  fine,  and  are  surrounded  by  the 
myelin  sheath,  but  do  not  possess  the  ordinary  external  nerve  sheath.  As 
they  pass  into  the  retina  they  lose  their  myelin  sheaths  and  proceed  as 
axis-cylinders  (the  cells  of  origin  of  these  fibers  are  in  the  retina).  Neuroglia 
supports  the  nerve  fibers  in  the  optic  nerve  trunk.  In  the  center  of  the  nerve 
is  a  small  artery,  the  arteria  centralis  retina.  The  number  of  fibers  in  the 


720 


THE    SENSES 


optic  nerve  is  said  to  be  upward  of  500,000  in  each  nerve  (Salzer).  The 
fibers  of  the  optic  nerve  spread  out  over  the  inner  surface  of  the  retina  as 
far  as  the  ora  serrata. 

The  retina  itself  consists  of  layers  of  nerve  elements  supported  by  deli- 
cate connective  tissue.  The  older  descriptions  recognize  some  eight  or  ten 
layers  in  the  retina.  These  appear  in  the  ordinary  microscopic  prepara- 
tions and  are  shown  in  figure  446.  But  the  newer  investigations  of  Cajal, 
Golgi,  Retzius,  and  others  have  shown  that  the  retina  is  a  much  simpler 
structure  than  heretofore  described.  The  retina  is  formed  of  essentially 
three  layers  of  nerve  and  sense  cells.  Naming  from  the  center  of  the  eye 
outward,  they  are:  The  ganglionic  layer;  the  layer  of  bipolar  cells;  and  the 
layer  of  rods  and  cones,  figure  447.  The  cells  of  these  layers  have  numer- 
ous fibrous  processes  which  interlock  in  such  a  way  that  they  seem  to  form 
different  areas  when  studied  in  cross-section,  figure  446.  If  we  recognize 
the  strata  of  interlacing  fibers,  then  the  following  may  be  made  out : 


The  layer  of  ganglion  cells  . . . 
The  layer  of  bipolar  cells    .... 


The  layer  of  visual  cells 


1.  Ganglionic  layer,  with  the  fibers  of  the 
optic  nerve. 

2.  Internal  molecular  layer. 

3.  Internal  nuclear  layer. 

4.  The  external  molecular  layer. 

5.  The  external  nuclear  layer. 

6.  The  layer  of  rods  and  cones. 




FIG.  447. — Transverse  Section  of  a  Mammalian  Retina.  A,  Layer  of  rods  and  cones; 
B,  bodies  of  visual  cells  (external  granular);  C,  external  molecular  layer;  E,  layer  of  bipolar 
cells  (internal  granular);  Ft  internal  molecular  layer;  G,  layer  of  ganglionic  cells;  H,  layer 
of  optic-nerve  fibers;  a,  rod;  b,  cone;  c,  body  of  the  cone  cell;  d,  body  of  the  rod  cell;  e, 
bipolar  rod  cells;/,  bipolar  cone  cells;  g,  h,  i,  j,  k,  ganglionic  cells  ramifying  in  the  various 
strata  of  the  internal  molecular  zone;  r,  inferior  arborization  of  the  bipolar  rod  cells,  con- 
necting with  the  ganglionic  cells;  r,  inferior  arborization  of  the  bipolar  cone  cells;  t,  epithelial 
or  Miiller  cells;  x,  point  of  contact  between  the  rods  and  their  bipolar  cells;  z,  point  of  con- 
tact between  the  cones  and  their  bipolar  cells;  s,  centrifugal  nerve  fiber.  (Cajal.) 


STRUCTURE    OF    THE    RETINA 


721 


The  Nerve  Fiber  and  Nerve  Cell  Layers. — The  inner  surface  of  the  retina 
is  formed  of  a  layer  of  nerve  fibers  which  have  their  origin  in  the  adjacent 
large  nerve  cells  and  converge  toward  the  exit  of  the  optic  nerve.  Extern- 
ally the  ganglionic  cells  send  up  numerous  processes,  or  dendrites,  which 
interlace  with  the  fibers  of  the  bipolar  cells  of  the  third  or  inner  nuclear 
layer  and  form  the  inner  molecular  layer  of  the  older  writers. 


FIG.  448. — Perpendicular  Section  cu  tne  Retina  of  a  Mammal.  At  External  grains  or 
bodies  of  rods;  B,  bodies  of  cones;  a,  horizontal  external  or  small  cell;  b,  horizontal  internal 
or  large  cell;  c,  horizontal  internal  cell  with  descending  protoplasmic  appendages;  e, 
flattened  arborization  of  one  of  the  large  cells;/,  g,  h,  j,  l>  spongioblasts  ramifying  in  the 
various  strata  of  the  internal  molecular  zone;  m,  n,  diffuse  spongioblasts;  o,  ganglionic  cell; 
i,  external  molecular  zone;  2,  internal  molecular  zone.  (Cajal.) 

The  Middle  Layer. — The  middle  layer  consists  of  bipolar  cells  which  send 
one  process  toward  the  ganglionic  layer  to  interlace  with  the  dendrites  of  the 
ganglionic  cells,  and  one  process  externally.  This  process  is  often  divided 
into  many  branches,  which  separate  out  into  a  horizontal  brush,  interlacing 


FIG.  449. — Distribution  of  the  Rods  and  Cones.     A,  In  the  peripheral  part  of  the  retina; 
B,  from  the  region  of  the  macula  lutea. 

with  the  processes  of  the  rods  and  cones.     Special  cells  have  been  described 
for  this  layer  of  the  retina,  as,  for  example,  the  spongioblasts  of  Cajal. 

The  External  Layer  of  Rods  and  Cones. — The  rod  cells  are  composed  of 
two  parts  quite  different  in  structure,  known  as  the  outer  and  inner  limbs. 

46 


FIG.  450. — Diagrammatic  Section  of 
the  Macula  Lutea  and  Fovea  Centralis, 
2,  Layer  of  nerve  fibers;  3,  layer  of 
multipolar  cells;  4,  internal  molecular 
layer,  composed  of  intertwining  arbor- 
escent processes;  5,  layer  of  bipolar 
cells,  or  internal  granular  layer;  6, 
external  molecular  layer,  composed 
of  intertwining  arborescent  processes; 
7,  nuclei  of  epithelial  cells,  or  external 
granular  layer;  8,  frillwork  formed 
by  processes  from  fibers  of  Miiller, 
often  called  the  "external  limiting 
membrane";  9,  layer  of  rods  and  cones; 
to,  layer  of  pigment  epithelium. 


SENSES 

The  outer  limb  is  a  cylindrical  rod 
about  30^  long  by  2/1  in  diameter.  It 
is  transparent  and  composed  of 
doubly  refractive  material.  The 
inner  limb  of  the  cell  is  about  the 
same  length  as  the  outer,  is  similar, 
and  is  longitudinally  striated,  and 
contains  a  nucleus  on  its  course, 
figure  447,  d. 

The  cone  cells  are  also  made  up 
of  two  limbs,  the  outer  of  which  is 
conical  instead  of  cylindrical  as  in 
the  case  of  the  rods.  In  other  re- 
spects they  are  similar  to  the  rods  in 
structure,  with  the  exception  that 
the  inner  limb  ends  in  a  brush  of 
fibrils  which  interlace  with  the  bipolar 
cells  of  the  middle  layer.  In  man 
and  mammals  the  number  of  rod 
cells  is  much  greater  than  the  cones, 
but  it  is  said  that  in  birds  cones  pre- 
dominate. Even  in  man  the  center  of 
the  fovea  centralis  is  devoid  of  rods 
and  consists  of  cones  only,  figure  450. 

All  the  elements  of  the  retina 
are  sustained  and  isolated  by  large 
cells  lying  vertically  which  are  known 
as  the  fibers  of  Miiller.  The  nucleus 
of  the  fiber  of  Miiller  is  found  at  the 
level  of  the  layer  of  bipolar  cells. 
The  two  extremities  of  the  proto- 
plasm or  cell  body  are  condensed  in 
two  homogeneous  layers,  known  as 
the  external  and  the  internal  limiting 
layer.  The  external  limiting  layer  is 
placed  just  between  the  two  segments 
of  the  rod  and  cone  cells,  forming  a 
fenestrated  membrane.  The  internal 
limiting  layer  is  situated  upon  the 
internal  surface  of  the  retina. 

At  the  ora  serrata  the  highly 
specialized  structure  of  the  retina 
disappears.  The  nerve  fibers  and 
ganglion  cells  disappear,  the  con- 
necting cells  are  fewer,  the  cones 
more  sparse,  and  the  rods  shorten 


STRUCTURE    OF    THE    RETINA 


723 


and   disappear.      The    structure    is  quickly  reduced  to  that  of  a  simple 
epithelial  membrane  known  as  the  pars  ciliaris  retinae. 

At  the  pars  ciliaris  retinae,  the  retina  is  represented  by  a  layer  of  columnar 


FIG.  451. — Diagram  of  the  Blood  Vessels  of  the  Human  Retina.  (Leber,  after  Jaeger.) 
arts,  vns,  Superior  nasal  artery  and  vein;  ats,  vts,  superior  temporal  artery  and  vein;  ani, 
uni,  inferior  nasal  artery  and  vein;  ati,  vti,  inferior  temporal  artery  and  vein;  ant,  vm, 
macular  artery  and  vein;  ane,  vme,  median  artery  and  vein. 

cells,  derived  from  the  fusion  of  the  nuclear  layers  which  are  in  contact  with 
the  pigment  layers  of  the  retina  and  continued  over  the  ciliary  processes. 
Pigment  Layer. — This  layer,  which  was  formerly  considered  part  of  the 


FIG.  452, — Blood  Vessels  of  the  Macula  Lutea.     The  part  that  is  totally  free  from  vessels  is 

the  fovea  centralis. 

choroid,  consists  of  cells  which  cover  and  entirely  surround  the  outer  limbs 
of  the  rods  and  cones. 

Blood  Vessels  of  the  Eyeball. — The  eye  is  very  richly  supplied  with  blood 
vessels.     In  addition  to  the  conjunctival  vessels,  which  are  derived  from  the 


724  THE    SENSES 

palpebral  and  lachrymal  arteries,  there  are  at  least  two  other  distinct  sets 
of  vessels  supplying  the  tunics  of  the  eyeball:  i,  the  vessels  of  the  sclera, 
choroid,  and  iris,  and  2,  the  vessels  of  the  retina.  The  first  are  the  short  and 
long  posterior  ciliary  arteries  which  pierce  the  sclerotic  in  the  posterior  half 
of  the  eyeball,  and  the  anterior  ciliary  which  enter  near  the  insertions  of  the 
recti.  These  vessels  anastomose  and  form  a  very  rich  choroidal  plexus; 
they  also  supply  the  iris  and  ciliary  processes,  forming  a  very  highly  vas- 
cular circle  round  the  outer  margin  of  the  iris  and  adjoining  portion  of 
the  sclerotic.  The  distinctness  of  these  vessels  from  those  of  the  conjunctiva 
is  well  seen  in  the  difference  between  the  bright  red  of  blood-shot  eyes 
(conjunctival  congestion),  and  the  pink  zone  surrounding  the  cornea  which 
indicates  deep-seated  ciliary  congestion. 

The  central  artery  of  the  optic  nerve  enters  the  retina  from  the  center  of 
the  optic  disc  and  sends  out  branches  over  the  retinal  cup  lying  in  the  nerve 
fiber  layer,  figure  451.  These  blood  vessels,  however,  are  absent  from  the 
fovea  centralis  and  reduced  in  size  in  the  macula  lutea,  figures  451  and  452. 

THE  OPTICAL  APPARATUS. 

The  optical  apparatus  may  be  supposed,  for  the  sake  of  description,  to 
consist  of  several  parts:  i,  a  system  of  transparent  refracting  surfaces  and 
media  by  means  of  which  images  of  external  objects  are  brought  to  a  focus 
upon  the  back  of  the  eye;  2,  a  sensitive  screen,  the  retina,  which  is  a  special- 
ized sensory  apparatus  in  connection  with  the  terminations  of  the  optic 
nerve,  and  capable  of  being  stimulated  by  luminous  objects,  and  of  sending 
such  impressions  as  to  produce  in  the  brain  visual  sensations.  To  these 
main  parts  may  be  added,  3,  an  apparatus  for  focusing  light  from  objects  at 
different  distances  from  the  eye,  and  4,  since  both  eyes  are  usually  employed 
in  vision,  an  arrangement  of  muscles  by  means  of  which  the  eyes  may  be 
turned  in  the  same  direction  so  that  binocular  vision  is  possible.  The 
arrangement  of  the  optic  nerve  fibers,  and  of  the  continuation  of  these  back- 
ward in  the  optic  chiasma,  and  thence  to  special  districts  of  the  brain 
have  already  been  discussed,  page  620. 

The  eye  may  be  compared  to  a  photographic  camera,  in  which  the  trans- 
parent refracting  media  correspond  to  the  photographic  lens.  In  a  camera 
images  of  external  objects  are  thrown  upon  a  screen,  the  sensitive  plate,  at 
the  back  of  the  camera  box.  In  the  eye,  the  camera  proper  is  represented 
by  the  eyeball  with  its  choroidal  pigment,  the  sensitive  screen  by  the  retina, 
and  the  lens  by  the  refracting  media.  In  the  case  of  the  camera,  the  screen 
is  adjusted  to  receive  clear  images  of  objects  at  different  distances  by  a 
mechanical  apparatus  for  focusing.  The  corresponding  adjustment  in  the 
eye  is  accomplished  by  the  accommodating  mechanism. 

Refractive  Media  and  Surfaces. — At  first  sight  it  would  s^em  as  if 
the  refracting  apparatus  of  the  eye  were  very  complicated,  since  it  consists 


IMAGE    FORMATION  725 

of  so  many  parts.  These  parts  are:  the  anterior  surface  of  the  cornea  itself, 
the  posterior  surface  of  the  cornea,  the  aqueous  humor,  the  anterior  surface 
of  the  lens,  the  substance  of  the  lens  itself  (which  is  unequally  refractive), 
the  posterior  surface  of  the  lens,  and  the  vitreous  humor.  Thus  there  are 
four  surfaces,  and  at  least,  including  the  air,  five  media.  For  all  practical 
purposes,  however,  we  may  leave  out  of  consideration  all  but  three  refracting 
surfaces  and  their  adjacent  media.  These  are:  the  anterior  surface  of  the 
cornea,  separating  the  air  and  the  corneal  substance;  the  anterior  surface 
of  the  lens,  separating  the  aqueous  humor  and  the  lens  substance;  and  the 
posterior  surface  of  the  lens,  separating  the  lens  surface  from  the  vitreous 
humor. 

Image  Formation. — In  the  refraction  through  a  simple  transparent 
spherical  surface  there  are  certain  cardinal  points  to  be  considered.  The 
rays  of  light  which  fall  perpendicularly  on  such  a  surface  pass  through  with- 
out refraction.  All  such  rays  cut  the  center  of  the  radius  of  curvature  of  the 


FIG.  453. — Diagram  of  a  Simple  Optical  System.  The  curved  surface,  bd,  is  supposed 
to  separate  a  less  refractive  medium  toward  the  left  from  a  more  refractive  medium 
toward  the  right.  N,  the  center  of  curvature  or  nodal  point;  OA,  principal  axis;  P, 
principal  point;  ab  and  OP,  rays  from  an  infinite  distant  point;  bN  and  dN,  secondary 
axes;  F  and  F2,  posterior  and  anterior  principal  foci;  PA  and  df",  parallel  rays  that 
meet  in  an  infinite  distance  only. 

lens,  called  the  nodal  point.  A  line  OA  that  passes  through  the  center  of 
curvature  of  a  lens  and  thus  pierces  the  nodal  point  N  is  called  the  optical 
axis,  and  the  point  on  the  surface  pierced  by  the  optical  axisP  is  the  princi~ 
pal  point.  In  every  optical  system  there  are  certain  other  cardinal  facts 
to  be  considered.  All  rays  which  do  not  strike  vertical  to  the  curved  sur- 
face are  refracted  toward  the  optical  axis.  Rays  which  impinge  upon  the 
spherical  surface  of  a  lens  parallel  to  the  optical  axis  will  meet  at  a  point 
upon  the  axis  called  the  posterior  principal  focus,  figure  453,  F.  The  pos- 
terior principal  focus  is  outside  of  the  nodal  point.  Again,  there  is  a  point 
in  the  optical  axis  in  front  of  the  surface,  rays  of  light  from  which  strike 
the  surface  so  that  they  are  refracted  in  a  line  parallel  with  the  axis, 
FtfLf";  such  a  point,  F%,  is  called  the  anterior  principal  focus. 
.  In  any  given  system  the  principal  foci  can  be  found  by  erecting  verticals 
at  the  nodal  and  principal  points  of  the  optical  axis  and  laying  off  lengths 
on  each,  a  and  b,  proportional  to  the  refractive  indices  of  the  media.  A  line 


726 


THE    SENSES 


drawn  through  a  on  the  principal  vertical  and  b  on  the  nodal  vertical  will  cut 
the  optical  axis  at  the  posterior  principal  focus,  and  vice  versa. 

If  a  luminous  point  outside  the  anterior  principal  focus  is  considered, 
it  is  obvious  that  rays  from  it  will  be  so  refracted  when  they  enter  the  convex 
surface  that  they  will  become  converging  and  will  ultimately  meet  again  in 
a  point  or  focus.  Two  such  points  form  conjugate  foci,  figure  454.  If  the 
anterior  focus  of  a  conjugate  is  moved  away  from  the  anterior  principal  focus, 
then  the  posterior  conjugate  will  move  toward  the  posterior  principal  focus, 
and  the  converse.  If  one  conjugate  is  known,  the  other  can  be  found  as 


FlG.  454. — Diagram  to  Show  the  Relations  of  Conjugate  Foci,     cd,y  Refracting  surface; 
AB  and  ba,  conjugate  foci;  o,  nodal  point;  F",  posterior  principal  focus. 

follows:  From  a  point  in  the  plane  of  the  known  conjugate,  but  outside  the 
principal  axis,  draw  two  rays,  one  perpendicular  to  the  refracting  surface 
which  will  pass  through  the  nodal  point,  the  other  parallel  to  the  principal 
axis.  The  latter  will  be  refracted  through  the  posterior  principal  focus  and 
when  prolonged  will  meet  the  first  ray  in  the  plane  of  the  second  conjugate, 
figure  454,  a.  This  relationship  between  conjugate  foci  is  played  upon  in  the 
focusing  of  a  camera. 

It  is  quite  obvious  that  the  eye,  even  considering  only  the  three  surfaces 
above  indicated,  is  a  much  more  complicated  optical  apparatus  than  the  one 
described  in  the  figure.  It  is,  however,  possible  to  reduce  the  refractive 
surfaces  and  media  to  a  simpler  form  when  the  refractive  indices  of  the  dif- 
ferent media  and  the  curvature  of  each  surface  are  known.  All  of  these 
data  have  been  very  carefully  collected.  They  are  as  follows: 

Index  of  refraction  of  aqueous  and  vitreous =    1.3365 

Index  of  refraction  of  the  lens =    i  .4371 

Radius  of  curvature  of  cornea =    7  . 829  mm. 

Radius  of  curvature  of  anterior  surface  of  lens =10.0       mm. 

Radius  of  curvature  of  posterior  surface  of  lens =    6.0       mm. 

Distance  between  anterior  surface  of  cornea  and  anterior  sur- 
face of  lens =  3.6  mm. 

Distance  between  anterior  surface  of  cornea  and  posterior 

surface  of  lens =  7.2  mm. 

With  these  data  it  has  been  found  comparatively  easy  by  mathematical 
calculation  to  reduce  the  different  refractive  surfaces  of  the  different  curva- 


IMAGE   FORMATION 


727 


tures  into  one  mean  curved  surface  of  known  curvature,  and  the  different 
refracting  media  into  one  mean  medium  the  refractive  power  of  which  is 
known. 

The  simplified  or  so-called  schematic  eye,  formed  upon  this  principle, 
suggested  by  Listing  as  the  reduced  eye,  has  the  following  dimensions: 

From  the  anterior  surface  of  the  cornea  to  the  principal  point  =    2  . 3448  mm. 

From  the  nodal  point  to  the  posterior  surface  of  lens =   o  .4764  mm. 

Posterior  principal  focus  lies  behind  cornea =22. 8237  mm. 

Anterior  principal  focus  in  front  of  cornea =  12  . 8326  mm. 

Radius  of  curvature  of  ideal  surface =    5. 1248  mm. 

In  this  reduced  or  simplified  eye  the  principal  posterior  focus,  about 
22.8  mm.  behind  the  spherical  surface,  would  correspond  to  the  position  of 
the  retina  behind  the  anterior  surface  of  the  cornea.  The  ideal  refracting 
surface  would  be  situated  about  midway  between  the  posterior  surface  of 
the  cornea  and  the  anterior  surface  of  the  lens. 

The  optical  axis  of  the  eye  is  a  line  drawn  through  the  centers  of  curva- 
ture of  the  cornea  and  lens,  and  when  prolonged  backward  it  cuts  the  retina 
between  the  optic  disc  and  fovea  centralis.  This  differs  somewhat  from 
the  visual  axis  which  passes  through  the  nodal  point  of  the  reduced  eye  to 


FIG.  455. — Diagram  of  the  Method  of  the  Formation  of  an  Inverted  Image  Exactly  Focused 
upon  the  Retina.     The  dotted  line  is  the  ideal  surface  of  curvature. 

the  fovea  centralis,  and  forms  an  angle  of  five  degrees  with  the  optical  axis. 
The  visual  or  optical  angle  is  the  angle  included  between  the  lines  drawn 
from  the  opposite  borders  of  any  object  through  the  nodal  point.  It  has 
been  shown  by  Helmholtz  that  the  smallest  angular  distance  between  two 
points  which  can  be  appreciated  is  fifty  seconds,  the  size  of  the  retinal  image 
being  3  . 65^.  This  practically  corresponds  to  the  diameter  of  the  cones  at 
the  fovea  centralis  which  is  3^,  the  distance  between  the  centers  of  two 
adjacent  cones  being  4/4. 

The  image  of  an  object  formed  upon  the  retina  may  be  considered  as  a 
series  of  points,  from  each  of  which  a  pencil  of  light  diverges  to  the  eye,  and 
this  pencil  has  for  its  center  or  axis  a  ray  which  impinging  upon  the  refract- 
ive surface  perpendicularly  to  the  surface  is  not  refracted^  but  passes 


7 2&  THE   SENSES 

through  the  nodal  point  and  is  prolonged  backward  to  the  retina.  The 
diverging  rays  are  refracted  to  converge  to  a  posterior  conjugate  focus 
behind  the  lens  on  the  chief  axis  of  the  pencil  of  light  proceeding  from  the 
point  in  question.  This  focus,  if  the  image  is  to  be  clear,  should  fall  on  the 
retina. 

Thus  from  each  point  of  an  object  a  corresponding  image  is  formed  on 
the  retina,  so  that  an  image  of  the  distant  object  is  produced.  It  is  an  inverted 
image.  Whether  the  image  is  blurred  or  not  depends  upon  the  refractive 
power  of  the  media  and  upon  the  distance  of  the  anterior  surface  of  the  cor- 


FIG.  456. — Diagram  of  the  Course  of  a  Ray  of  Light,  to  Show  how  a  Blurred  or 
Indistinct  Image  is  Formed  if  the  Object  be  not  Exactly  Focused  upon  Retina.  The 
surface  CC  should  be  supposed  to  represent  the  ideal  curvature.  The  nodal  point  should 
be  nearer  the  posterior  surface  of  lens  as  in  figure  455. 

nea  from  the  retina.  If  the  refractive  media  are  too  powerful  or  the  eye  too 
long,  the  image  is  formed  in  front  of  the  retina,  figure  456;  if  the  reverse,  the 
image  is  formed  behind  the  retina,  and  in  both  cases  an  indistinct  and  blurred 
image  is  the  result. 

Accommodation. — The  distinctness  of  the  image  formed  upon  the 
retina  is  mainly  dependent  on  the  perfection  with  which  the  rays  emitted 
by  each  luminous  point  of  the  object  are  brought  to  a  focus  upon  the  retina. 
If  this  focus  occurs  at  a  point  either  in  front  of  or  behind  the  retina,  indis- 
tinctness of  vision  ensues,  in  the  way  we  have  just  described,  with  the  pro- 
duction of  a  halo.  The  focal  distance,  i.e.,  the  distance  of  the  point  at  which 
the  luminous  rays  are  collected  from  a  lens,  besides  being  regulated 
by  the  degree  of  convexity  and  density  of  the  lens,  varies  with  the  distance 
of  the  object  from  the  lens,  being  greater  as  the  distance  is  shorter,  and  vice 
versa.  In  other  words,  the  luminous  points  on  the  object  and  the  focal  points 
on  the  retina  are  conjugate  foci.  Hence,  since  objects  placed  at  various 
distances  from  the  eye  can  within  certain  limits  be  seen  with  almost  equal 
distinctness,  there  must  be  some  provision  by  which  the  eye  is  enabled  to 
adapt  itself,  so  that,  at  whatever  distance  the  luminous  object  may  be,  the 
focal  point  may  always  fall  exactly  upon  the  retina. 

Accommodation  is  the  act  of  adapting  the  eye  to  vision  at  different  distances. 
It  is  obvious  that  the  effect  might  be  produced  in  either  of  two  ways,  viz., 


ACCOMMODATION  729 

i,  by  altering  the  convexity,  and  thus  the  refracting  power,  either  of  the  cornea 
or  of  the  lens;  or  2,  by  changing  the  position  of  the  lens  relative  to  the  retina, 
as  in  the  focusing  of  a  camera,  so  that  whether  the  object  be  near  or  distant, 
the  focal  points  to  which  the  rays  are  converged  by  the  lens  may  always  fall 
exactly  on  the  retina.  The  amount  of  either  of  these  changes  which  is  re- 
quired in  even  the  widest  range  of  vision  is  extremely  small,  for  from  the  re- 
fractive powers  of  the  media  of  the  eye  the 
difference  between  the  focal  distances  of  the 
images  of  an  object  at  a  distance  and  of  one 
at  four  inches  is  only  about  3 . 5  mm.  On 
this  calculation  the  change  in  the  distance  of 
the  retina  from  the  lens  required  for  vision  at 
all  distances,  supposing  the  cornea  and  lens 
to  remain  the  same,  would  not  be  more  than 

about  2.5   mm.      Beer  has  shown  that  the 

,      ....      ,     ,  FIG.  4157. — Diagram  Showing 

second  method  is  indeed  the  type  of  accom-  Three  Reflections  of  a  Candle. 

modative  apparatus  in  fishes.     But  in  man  i,  From  the  anterior  surface  of 

.                .       i                              .  cornea;    2,    from    the   antenor 

and  the  higher  animals  accommodation  occurs  surface  of  lens;  3,  from  the 

by  the  first  method,  i.e..  by  changing  the  con-  posterior  surface  of  lens.    For 

.    .  further   explanation,    see   text. 

vexity  of  the  refracting  surface.  The  experiment  is  best  per- 

The  accommodation  of  the  human  eye  for  formed  by  employing  an  in- 

.         .         MI  strument   invented    by    Helm- 

objects  at  different  distances  is  primarily  due  holtz,  termed  a  Phakoscope. 

to  a  varying  shape  of  the  lens,  its  front  surface 

becoming  more  or  less  convex,  according  as  the  distance  of  the  object  looked 
at  is  near  or  far.  The  nearer  the  object,  the  more  convex  the  front  surface 
of  the  lens,  up  to  a  certain  limit,  and  vice  versa;  the  back  surface  takes  little 
or  no  share  in  accommodation.  The  following  simple  experiment  illustrates 
this  point:  If  a  lighted  candle  be  held  a  little  to  one  side  of  a  person's  eye, 
an  observer  looking  at  the  eye  from  the  other  side  sees  three  distinct  images 
of  the  flame,  figure  457.  The  first  and  brightest,  i,  is  a  small  erect  image 
formed  by  the  anterior  convex  surface  of  the  cornea;  the  second,  2,  is  also  erect, 
but  larger  and  less  distinct  than  the  preceding,  and  is  formed  at  the  anterior 
convex  surface  of  the  lens;  the  third,  3,  is  smaller,  inverted,  and  indistinct; 
it  is  formed  at  the  posterior  surface  of  the  lens,  which  is  concave  forward,  and 
therefore,  like  all  concave  mirrors,  gives  an  inverted  image.  If  the  eye 
under  observation  be  made  to  look  at  a  near  object,  the  second  image  be- 
comes smaller,  clearer,  and  approaches  the  first.  If  the  eye  be  now  adjusted 
for  a  far  point,  the  second  image  enlarges  again,  becomes  less  distinct,  and 
recedes  from  the  first.  In  both  cases  alike  the  first  and  third  images  remain 
unaltered  in  size,  distinctness,  and  relative  position.  This  proves  that  during 
accommodation  for  near  objects  the  curvature  of  the  cornea,  and  that  of  the 
posterior  surface  of  the  lens,  remain  unaltered,  while  the  anterior  surface  of 
the  lens  becomes  more  convex  and  approaches  the  cornea. 


THE   SENSES 

The  experiment,  figure  458,  is  more  striking  when  the  two  prisms  of  the 
phakoscope  which  form  two  images  of  the  candle  are  used.  The  pair  of 
images  of  the  candle  from  the  front  surface  of  the  lens  not  only  approach 
those  from  the  cornea  during  accommodation,  but  also  approach  one  another, 
and  become  somewhat  smaller,  Sanson's  images. 


FIG.  458. — Diagram  of  Sanson's  Images.  A,  When  the  eyes  are  focused  for  far 
objects,  and  B,  when  they  are  focused  for  near  objects.  The  figure  to  the  right  in  A  and 
B  is  the  inverted  image  from  the  posterior  surface  of  the  lens. 


\Cire.M. 

Had.M  SupACerv.C, 


FIG.  459. — Diagrammatic  representation  of  the  nerves  of  the  intrinsic  muscles  of  the 
eye.  Sup.  Corp.  Quad.,  superior  corpora  quadrigemina.  Nuc.  Ill,  nucleus  of  the  third 
cranial  nerve.  Sup.  Cerv.  G.,  superior  cervical  ganglion.  Circ.  M .,  circular  muscles  of 
the  iris.  Rod.  M.,  radial  muscles  of  the  iris.  Ciliary  G.,  ciliary  ganglion  (Greene). 


RANGE    OF  DISTINCT  VISION  731 

The  Mechanism  of  Accommodation. — The  mechanism  of  accommo- 
dation depends  primarily  upon  the  inherent  tendency  of  the  lens  to  approxi- 
mate the  shape  of  a  sphere.  When  the  eye  is  at  rest  the  intra-ocular  tension 
is  such  as  to  put  stress  on  the  suspensory  ligament  around  its  equator,  which 
compresses  the  elastic  lens  in  its  antero-posterior  dimension.  The  elasticity 
of  the  lens  can  make  itself  apparent  when  the  tension  of  the  suspensory  liga- 
ment is  relaxed.  This  takes  place  completely  after  a  division  of  the  fibers 
of  the  zonula.  When  we  remove  the  lens  from  the  eye  of  a  young  person, 
we  see  it  assume  the  spherical  shape  immediately  upon  the  division  of  its 
capsule.  In  life  the  slackening  of  the  tension  of  the  suspensory  ligament 
of  the  lens  is  brought  about  by  the  active  contractions  of  the  muscle  fibers 
of  the  ciliary  body,  the  combined  contractions  of  the  radial  and  the  circu- 
lar fibers.  This  allows  the  surfaces  of  the  lens  by  its  own  elastic  powers  to 
become  more  convex,  thus  focusing  entering  rays  of  light  from  a  nearer 
object  upon  the  retina,  figure  460.  It  therefore  appears  that  when  the  eye 


FIG.  460. — Diagram  Representing  by  Dotted  Lines  the  Alteration  in  the  Shape  of  the  Lens 
on  Accommodation  for  Near  Objects.     (E.  Landolt.) 

is  at  rest  it  is  focused  for  distant  objects,  in  as  much  as  the  suspensory 
ligament  is  taut  and  the  anterior  surface  of  the  lens  more  flattened.  The 
normal  eye  is  therefore  passive  when  in  focus  for  distant  objects.  It  is 
the  active  contraction  of  the  muscles  of  accommodation  that  focuses  for 
near  objects. 

The  iris  acts  in  co-ordination  with  the  accommodative  contractions  of 
the  ciliary  muscles.  In  viewing  near  objects  the  pupil  contracts,  and  upon 
dewing  distant  ones  it  dilates. 

Range  of  Distinct  Vision. — Near-point. — In  every  eye  there  is  a  limit  to 
the  power  of  accommodation.  If  a  book  be  brought  nearer  and  nearer 
to  the  eye,  the  type  at  last  becomes  indistinct,  and  cannot  be  brought  into 
focus  by  any  effort  of  accommodation,  however  strong.  The  printed 
letters  appear  gray  and  with  shadowy  outlines.  At  a  certain  distance  the 


732 


THE    SENSES 


letters  are  black  and  distinct  but  just  within  this  point  the  outlines  present 
a  just  perceptible  indistinctness,  a  loss  of  sharpness  of  border.  This  limit 
is  termed  the  near -point  of  vision.  The  near-point  can  be  determined 
by  the  experiment  of  Scheiner.  Two  small  holes  not  more  than  2  mm. 
apart  are  pricked  in  a  card  with  a  pin;  at  any  rate  their  distance  from 
each  other  must  not  exceed  the  diameter  of  the  pupil.  The  card  is  held 
with  the  holes  close  in  front  of  the  eye,  and  a  small  needle  viewed 
through  the  pin-holes.  At  a  moderate  distance  it  can  be  clearly  focused, 
but  when  brought  nearer,  within  a  certain  point,  the  image  appears 
double  and  more  or  less  blurred.  This  point  where  the  needle  ceases 
to  appear  single  is  the  near-point  of  vision.  Its  distance  from  the  eye 
can  of  course  be  readily  measured.  It  is  usually  about  five  or  six  inches, 
12  to  15  cm.  In  the  accompanying  figure,  461,  the  lens  b  represents  the 
eve;  e>  />  tne  two  pin-holes  in  the  card;  nn  the  retina;  a  represents 


FIG.  461. — Diagram  of  Experiment  to  Ascertain  the  Minimum  Distance  of  Distinct  Vision. 

the  position  of  the  needle.  When  the  needle  is  at  a  moderate  distance, 
the  two  pencils  of  light  coming  through  the  holes  e  and  /  are  focused  at  a 
single  point  on  the  retina  nn.  If  the  needle  be  brought  nearer  than  the 
near.-point,  the  strongest  effort  of  accommodation  is  not  sufficient  to  focus 
the  two  pencils,  they  meet  at  a  point  behind  the  retina.  The  effect  is  the 
same  as  if  the  retina  were  shifted  forward  to  mm.  Two  indistinct  images, 
h,  g,  are  formed  by  the  converging  pencils  of  light,  one  from  each  hole. 
It  is  interesting  to  note  that  when  in  this  way  two  shadowy  images  are 
produced,  the  lower  one,  g,  really  appears  out  in  space  in  the  position  Q, 
while  the  upper  one  appears  in  the  position  P.  This  may  be  readily  veri- 
fied by  covering  the  holes  in  succession.  This  is  due  to  the  fact  that  when 
points  on  the  retina  outside  the  visual  axis  are  stimulated  the  sensation  is 
referred  to  an  object  in  space  along  the  line  of  the  secondary  optic  axis 
that  cuts  the  retina  at  the  point  stimulated. 

During  accommodation  two  other  changes  take  place  in  the  eyes.  The 
two  eyes  converge  by  the  action  of  the  extra-ocular  muscles,  chiefly  by  the 
internal  and  inferior  recti  or  internal  and  superior  recti.  The  pupils  also 
contract. 


REFLEXES    OF    THE  PUPIL 


733 


Movements  of  the  Eye.  The  eyeball  possesses  movement  around  three  axes 
indicated  in  figure  462,  viz.,  an  antero-posterior,  a  vertical,  and  a  transverse, 
passing  through  a  center  of  rotation  a  little  behind  the  centre  of  the  optic  axis. 
The  movements  are  accomplished  by  pairs  of  muscles. 


Direction  of  Movement. 


Inward . . 
Outward . 


Upward 


Downward 


Inward  and  upward 


Inward  and  downward. 


Outward  and  upward. 


Outward  and  downward. 


By  what  muscles  accomplished. 

Internal  rectus. 

External  rectus. 
/  Superior  rectus. 
1  Inferior  oblique. 
/  Inferior  rectus. 

(Superior  oblique. 
Internal  and  superior  rectus. 
!    Inferior  oblique. 
Internal   and   inferior  rectus. 
Superior  oblique. 
(    External  and  superior  rectus. 
'     Inferior  oblique, 
f    External  and  inferior  rectus. 
\    Superior  oblique. 


FIG.  462. — Diagram  of  the  Axes  of  Rotation  of  the  Eye.     The  thin  lines  indicate  axes  of 
rotation,  the  thick  the  position  of  muscular  attachment. 

The  contraction  of  all  of  the  muscles  during  the  act  of  accommodation, 
viz.,  of  the  ciliary  muscles,  of  the  recti  muscles,  and  of  the  sphincter  pupillae, 
is  under  the  control  of  the  fibers  of  the  third  nerve.  But  the  superior  oblique 
may  also  be  employed,  in  which  case  the  fourth  nerve  is  concerned. 

Reflexes  of  the  Pupil. — Contraction  of  the  iris  may  occur  under  the 
following  circumstances:  i.  On  exposure  of  the  eye  to  a  bright  light.  On 


734  THE    SENSES 

the  local  application  of  eserine  (active  principle  of  Calabar  bean).  3.  On  the 
administration  internally  of  opium,  aconite,  and  in  the  early  stages  of  chloro- 
form and  alcohol  poisoning.  4.  On  division  of  the  cervical  sympathetic  or  of 
stimulation  of  the  third  nerve.  Dilatation  of  the  pupil  occurs,  i,  in  a  dim 
light;  2,  when  the  eye  is  focused  for  distant  objects;  3,  on  the  local  applica- 
tion of  atropine  and  its  allied  alkaloids;  4,  on  the  internal  administration  of 
atropine  and  its  allies;  5,  in  the  later  stages  of  poisoning  by  chloroform, 
ether,  and  other  drugs;  6,  on  paralysis  of  the  third  nerve;  7,  on  stimulation 
of  the  cervical  sympathetic,  or  of  its  center  in  the  floor  of  the  front  of  the 
aqueduct  of  Sylvius.  The  contraction  of  the  pupil  is  under  the  control  of 
a  center  in  the  floor  of  the  aqueduct  beneath  the  anterior  corpora  quadri- 
gemina.  This  center  is  reflexly  stimulated  by  a  bright  light,  and  the  dilata- 
tion when  the  center  is  not  in  action  is  due  to  the  stimulation  of  the  radial 
fibers  of  the  iris  by  sympathetic  nerves.  In  addition,  it  appears  that  both 
contraction  and  dilatation  may  be  produced  by  a  local  action  of  certain  drugs 
which  is  independent  of,  and  probably  often  antagonistic  to,  the  action  of  the 
central  apparatus  of  the  third  and  sympathetic  nerves. 

The  close  co-ordination  between  the  two  eyes  is  nowhere  better  shown 
than  by  the  condition  of  the  pupil.  If  one  eye  be  shaded  by  the  hand  its 
pupil  will  of  course  dilate;  the  pupil  of  the  other  eye  will  also  dilate,  though 
unshaded,  due  to  crossed  reflex  action. 

Defects  in  the  Optical  Apparatus. — Under  this  head  we  may  con- 
sider the  defects  known  as:  i,  Spherical  Aberration;  2,  Chromatic  Aberra- 
tion; 3,  Astigmatism;  4,  Myopia;  5,  Hypermetropia. 

The  normal  or  emmetropic  eye  is  so  perfect  that  parallel  rays  are  brought 
exactly  to  a  focus  on  the  retina  without  any  effort  of  accommodation,  figure 
466.  Hence  all  objects  except  near  ones  (in  practice  all  objects  at  a  distance 
of  twenty  feet  or  more)  are  seen  without  any  effort  of  accommodation;  in 
other  words,  the  far-point  of  the  normal  eye  at  rest  is  at  an  infinite  distance. 
In  viewing  near  objects  we  are  conscious  of  the  effort  (the  contraction  of 
the  ciliary  muscle)  by  which  the  anterior  surface  of  the  lens  is  rendered 
more  convex,  and  rays  which  would  otherwise  be  focused  behind  the  retina 
are  converged  upon  the  retina. 

Spherical  Aberration. — The  rays  in  a  cone  of  light  from  a  point  on  an 
object  situated  in  the  field  of  vision  do  not  all  meet  in  the  same  point  in 
the  retina,  owing  to  the  greater  refraction  of  the  rays  which  pass  through 
the  margin  of  a  lens  than  those  traversing  its  central  portion.  This  defect 
is  spherical  aberration.  In  the  camera,  telescope,  microscope,  and  other 
optical  instruments  it  is  remedied  by  the  interposition  of  a  screen  with  a 
circular  aperture  in  the  path  of  the  rays  of  light,  cutting  off  all  the  marginal 
rays  and  allowing  the  passage  only  of  those  near  the  center.  Such  cor- 
rection is  effected  in  the  eye  by  the  iris,  which  forms  a  diaphragm  to  cover  the 
circumference  of  the  lens,  and  prevents  the  rays  from  passing  through  any 


CHROMATIC  ABERRATION  735 

part  of  the  lens  but  its  center,  which  corresponds  to  the  pupil.  The  iris  is 
pigmented  to  prevent  the  passage  of  rays  of  light  through  its  substance. 
The  image  of  an  object  will  be  most  clearly  denned  and  distinct  when  the 
pupil  is  small,  if  the  light  is  abundant;  so  that,  while  a  sufficient  number 
of  rays  are  admitted,  the  narrowness  of  the  pupil  may  prevent  the  pro- 
duction of  indistinctness  of  the  image  by  spherical  aberration.  But  even 
the  image  formed  by  the  rays  passing  through  the  circumference  of  the 
lens,  when  the  pupil  is  much  dilated,  as  in  the  dark  or  in  a  feeble  light, 
may  be  well  denned.  Some  types  of  optical  apparatus  are  corrected  for 
this  defect  by  a  central  instead  of  marginal  diaphragm. 

Distinctness  of  vision  is  further  secured  by  the  pigment  of  the  outer  sur- 
face of  the  retina  and  of  the  posterior  surface  of  the  iris  and  the  ciliary  proc- 
esses. This  absorbs  any  rays  of  light  that  may  be  reflected  within  the  eye 
and  prevents  their  being  thrown  again  upon  the  retina  so  as  to  interfere 
with  the  images  formed  there.  The  pigment  of  the  retina  is  especially  im- 
portant in  this  respect;  for  with  the  exception  of  its  outer  layer  the  retina  is 
very  transparent;  and  if  the  surface  behind  it  were  not  of  a  dark  color,  but 
capable  of  reflecting  the  light,  the  luminous  rays  which  had  already  acted 
on  the  retina  would  be  reflected  again  and  would  fall  upon  other  parts  of 
the  same  membrane,  producing  indistinctness  of  the  images. 

Chromatic  Aberration. — In  the  passage  of  light  through  the  periphery 
of  an  ordinary  convex  lens  decomposition  of  each  ray  into  its  elementary 
colors  commonly  ensues,  and  a  colored  margin  appears  around  the  image, 
owing  to  the  unequal  refraction  which  the  elementary  colors  undergo. 
This  is  termed  chromatic  aberration.  It  is  corrected  by  the  use  of  lenses 
constructed  of  alternate  layers  of  glass  of  different  refractive  indices  so 
ground  that  they  produce  chromatic  dispersion  in  opposite  directions  and 
thus  mutually  correct  any  chromatic  aberration  which  may  have  resulted. 
The  human  eye  has  considerable  chromatic  aberration,  as  may  readily  be 
demonstrated,  Experiment  13,  page  721. 

An  ordinary  ray  of  white  light  in  passing  through  a  prism  has  its  con- 
stituent rays  refracted  in  unequal  degrees,  and  therefore  appears  as  colored 
bands  fading  off  into  each  other,  known  as  the  spectrum.  The  colors  of 
the  spectrum  are  arranged  as  follows;  red,  orange,  yellow,  green,  blue, 
indigo,  violet;  of  these  the  red  ray  is  the  least,  and  the  violet  the  most 
refracted.  Hence,  as  Helmholtz  has  shown,  the  rays  from  a  point  of  white 
light  cannot  be  accurately  focused  on  the  retina,  for  if  we  focus  for  the 
red  rays,  the  violet  are  out  of  focus,  and  vice  versa;  such  objects,  if  not  ex- 
actly focused,  are  often  seen  surrounded  by  a  pale  yellowish  or  bluish 
fringe. 

For  similar  reasons  a  red  surface  looks  nearer  than  a  blue  one  at  an 
equal  distance,  because,  the  red  rays  being  less  refrangible,  a  stronger 
effort  of  accommodation  is  necessary  to  focus  them,  and  the  eye  is  ad- 


73 6  THE   SENSES 

justed  as  if  for  a  nearer  object,  and  therefore  the  red  surface  appears 
nearer,  Experiment  13. 

Astigmatism. — The  formation  of  perfect  images  becomes  impossible 
when  either  of  the  refractive  surfaces  of  the  eye  has  unequal  curvatures  in 
the  different  meridians.  This  defect  is  called  astigmatism.  It  was  first 
discovered  by  Airy.  An  astigmatic  eye  cannot  form  a  perfect  image  of  a 
luminous  point,  and  since  images  of  objects  are  built  up  of  the  images  of 
the  infinite  number  of  points  on  their  surfaces  it  follows  that  all  images  are 
distorted.  Luminous  points  appear  with  imperfect  or  star-shaped  radii 
or  haloes.  If  these  radii  overlap  in  an  image  they  tend  to  neutralize  the 
diffuse  stimulating  effects  and  such  regions  appear  more  distinct,  while  in 
the  converse  condition  the  images  are  more  blurred,  figure  465. 

This  defect,  which  is  generally  present  in  a  slight  degree  in  all  eyes,  is 
usually  seated  in  the  cornea,  but  occasionally  in  the  lens  as  well. 

The  plane  of  greatest  curvature  in  the  cornea  is  usually  in  the  vertical 
meridian,  a  fact  which  doubtless  comes  from  the  pressure  of  the  eyelids 
during  development.  If  one  looks  at  figure  463,  A  or  B,  with  one  eye,  the 
three  iines  in  the  radii  of  the  figure  will  be  seen  with  unequal  distinctness. 
Certain  sets  will  stand  out  sharp  and  black  and  others  dim  and  with  indistinct 
outlines,  and  if  the  astigmatism  is  great  enough  the  three  lines  may  not  be 
distinguished.  Figures  C  and  D  of  this  series  enable  one  to  detect  minute 
traces  of  astigmatism  with  great  accuracy. 


ABC 

FIG.  463. — Astigmatic  Charts. 

It  is  somewhat  difficult  to  picture  the  rays  from  a  luminous  point  in  their 
course  through  eyes  which  have  this  defect,  but  an  examination  of  figure 
464  will  show  their  refraction  in  astigmatism.  In  this  figure  four  of  the  total 
sphere  of  rays  diverging  from  the  point  L  in  the  arrows  are  represented  as 
striking  on  the  refractive  surface  of  the  eye  a,tA,B,  C,  D,  and  being  converged 
toward  a  focus.  The  rays  A ,  C,  separated  by  a  vertical  line  on  the  refractive 
surface,  are  focused  at  flt  while  the  rays  A ,  B,  separated  by  the  horizontal 
distance  on  the  refractive  surface,  are  brought  to  a  focus  at/2.  Rays  from  the 
point  L,  therefore,  have  two  apparent  focal  points,  one  point  composed  of  the 
rays  that  strike  the  refractive  surfaces  in  a  horizontal  plane,  /2,  the  other  of 
rays  that  strike  in  a  vertical  plane,/.  If  the  retina  of  the  eye  be  placed  at/j, 
it  will  receive  an  image  of  a  luminous  point  with  indistinct  horizontal  halos 


MYOPIA 


737 


composed  of  the  unfocused  rays  of  all  other  meridians  than  the  vertical  which 
are  in  focus.  If  placed  at  the  position/2  it  will  receive  a  luminous  point  with 
indistinct  halos  in  the  vertical  plane.  If  the  series  of  points  in  the  arrow  MN 
be  considered,  it  is  evident  that  at  the  position/!  the  rays  which  fall  in  the  ver- 
tical plane  will  form  distinct  foci,  while  those  that  fall  in  the  horizontal  plane 
will  form  overlapping  diffuse  images  in  that  plane.  Since  they  are  over- 
lapping, they  will  not  appear  separate  except  at  the  ends  of  the  image  of  the 
arrow,  and  the  arrow  will  therefore  be  seen  distinctly.  If  the  position /2  is 


FIG.  464. — The  Unequal  Refraction  of  Rays  in  an  Astigmatic  Eye.     (John  Green.) 

considered  where  the  rays  of  the  horizontal  plane  are  focused,  then  it  is 
evident  that  the  points  in  the  arrow  MN  will  present  a  series  of  rays  or  halos 
in  the  vertical  plane,  thus  rendering  its  outline  very  dim  or  indistinct.  The 
condition  with  the  arrow  OP  is  exactly  the  reverse.  Hence,  in  the  astigmatic 
eye  the  images  of  the  horizontal  arrow  MN  will  be  distinct  at  the  focus/!, 
while  the  image  of  the  vertical  arrow  OP  will  be  distinct  in  the  focus /2,  and 
the  eye  cannot  see  the  two  lines  distinctly  at  the  same  time.  This  condition  is 
further  illustrated  in  figure  465  which  represents  the  type  of  image  formed  at 
the  position  f,  shown  in  figure  464. 


FIG.  465. — Diagram  of  Character  of  Retinal  Images  in  Astigmatism.     (John  Green.) 

Myopia. — This  is  that  refractive  condition  of  the  eye  in  which  parallel 
rays  are  brought  to  focus  in  front  of  the  retina  when  the  eye  is  at 
rest,  4,  figure  466.  It  is  due  either  to  an  abnormal  elongation  of  the 
eyeball  antero-posteriorly  or  to  an  increase  in  the  convexity  of  the  re- 
fracting surfaces,  or  to  both  of  these  conditions.  Parallel  rays  from  a 
distant  point  are  focused  in  front  of  the  retina,  and,  crossing,  form  circles 


738 


THE    SENSES 


on  the  retina.  Thus,  the  images  of  distant  objects  are  blurred  and  in- 
distinct. The  eye  is,  as  it  were,  permanently  adjusted  for  a  near  point. 
Rays  from  a  point  near  the  eye  are  exactly  focused  on  the  retina.  But 
those  which  issue  from  any  object  beyond  a  slight  distance,  the  myopic 
far-point,  which  is  less  than  twenty  feet,  cannot  be  distinctly  focused. 
This  defect  is  corrected  by  concave  glasses,  which  cause  parallel  rays  to 
diverge  before  entering  the  eye.  Such  glasses  of  course  are  needed  only 
to  give  a  clear  vision  of  distant  objects.  For  near  objects  they  are  not 
required. 


FIG.  466. — Diagram  Showing:  i,  Normal  or  emmetropic  eye  bringing  parallel  rays 
exactly  to  a  focus  on  the  retina;  2,  normal  eye  at  rest,  showing  that  light  from  a  near  point  is 
focused  behind  the  retina,  but  by  increasing  the  curvature  of  the  anterior  surface  of  the 
lens  (shown  by  dotted  lines)  the  rays  are  focused  on  the  retina;  3,  hypermetropic  eye.  In 
this  case  the  axis  of  the  eye  is  shorter  and  the  lens  normal  (or  the  lens  may  be  flatter  than 
normal  and  the  eyeball  normal) ;  parallel  rays  are  focused  behind  the  retina;  4,  myopic  eye. 
In  this  case  the  lens  is  too  convex  (or  the  axis  of  the  eye  is  abnormally  long);  parallel  rays 
are  focused  in  front  of  the  retina. 

Hypermetropia. — This  is  that  refractive  condition  of  the  resting  eye 
in  which  parallel  rays  are  still  converging  but  not  yet  focused  at  the  retina, 
are  brought  to  a  focus  behind  the  retina,  3,  figure  466.  It  is  the  opposite 


VISUAL    SENSATIONS,  FROM    EXCITATION    OF    THE   RETINA       739 

of  myopia,  and  is  due  either  to  an  abnormal  shortening  of  the  eyeball 
antero-posteriorly  or  to  a  decrease  in  the  convexity  of  the  refracting 
surfaces,  or  both.  Parallel  rays  entering  the  eye  at  rest  are  focused  be- 
hind the  retina.  An  effort  of  accommodation  is  therefore  required  to 
focus  parallel  rays  on  the  retina.  When  the  rays  are  sharply  divergent, 
as  in  viewing  a  very  near  object,  the  accommodation  is  insufficient  to 
focus  them.  Thus,  not  only  do  distant  objects,  normally  seen  without 
effort,  require  an  act  of  accommodation,  but  near  objects  are  focused 
only  by  the  maximal  muscular  contraction  of  the  accommodative  mech- 
anism. The  eye  is  under  a  constant  strain  which  produces  in  the  end 
various  nervous,  as  well  as  ocular,  disorders.  This  defect  is  obviated 
by  the  use  of  convex  glasses,  which  render  the  pencils  of  light  more  con- 
vergent. Such  glasses  are  especially  needed  for  near  objects,  as  in  read- 
ing, etc.  They  are  also  required  for  distant  vision  to  rest  the  eye  by 
relieving  the  ciliary  muscle  from  constant  work. 

Presbyopia. — Presbyopia  is  a  condition  of  diminished  range  of  ac- 
commodation. It  takes  place  with  considerable  uniformity  from  youth  to 
old  age.  It  is  not  a  disease,  but  a  physiological  process  which  every  eye 
undergoes  as  its  owner  grows  older.  It  is  due  to  a  gradual  diminution  of 
elasticity  of  the  lens  by  a  sort  of  sclerosis  from  the  center  toward  the 
periphery.  It  begins  even  in  childhood,  but  advances  so  slowly  that  it  is 
not  until  the  age  of  twenty-five  or  so  that  a  distinct,  though  small,  nucleus 
is  present.  With  advancing  years  the  process  goes  on  until  finally  the 
lens  becomes  inelastic  and  is  unable  to  assume  a  shape  convex  enough  to 
focus  rays  from  a  near  object  upon  the  retina,  as  in  reading.  The  defect 
is  remedied  by  the  use  of  convex  lenses  equivalent  to  the  loss  in  accom- 
modation. 

Visual  Sensations,  from  Excitation  of  the  Retina. — Light  is  the 
normal  agent  in  the  excitation  of  the  retina.  The  only  portion  of  the  retina 
capable  of  reacting  to  the  stimulus  is  the  rod  and  cone  layer.  The  proofs  of 
this  statement  may  be  summed  up  thus:  i.  The  point  of  entrance  of  the  optic 
nerve  into  the  retina,  where  the  rods  and  cones  are  absent,  is  insensitive  to 
light  and  is  called  the  blind  spot.  The  phenomenon  itself  is  very  readily 
demonstrated.  If  we  close  one  eye,  and  direct  the  other  upon  a  point  at 
such  a  distance  to  the  side  of  any  object  that  the  image  of  the  latter  must 
fall  upon  the  retina  at  the  point  of  entrance  of  the  optic  nerve,  its  image  is 
lost.  If,  for  example,  we  close  the  left  eye,  and  direct  the  axis  of  the  right 
eye  steadily  toward  the  circular  spot  in  figure  467,  while  the  page  is  held  at 
a  distance  of  about  six  inches  from  the  eye,  both  dot  and  cross  are  visible. 
On  gradually  increasing  the  distance  between  the  eye  and  the  object,  by 
removing  the  book  farther  and  farther  from  the  face,  keeping  the  right  eye 
steadily  on  the  dot,  it  will  be  found  that  suddenly  the  cross  disappears  from 
view,  while  on  removing  the  book  still  farther  it  suddenly  comes  into  view 


74°  THE    SENSES 

again.  The  cause  of  this  phenomenon  is  simply  that  the  portion  of  retina 
which  is  occupied  by  the  entrance  of  the  optic  nerve  is  quite  blind;  and  there- 
fore that  when  it  alone  occupies  the  field  of  vision  objects  cease  to  be  visible. 


FIG.  467.  —  Diagram  for  Demonstrating  the  Blind  Spot. 

2.  In  the  fovea  centralis  and  macula  lutea,  which  contain  rods  and  cones  but 
no  optic-nerve  fibers,  light  produces  the  greatest  effect.  In  the  latter,  cones 
occur  in  large  numbers,  and  in  the  former  cones  without  rods  are  found, 
whereas  in  the  rest  of  the  retina,  which  is  not  so  sensitive  to  light,  there  are 
fewer  cones  than  rods.  We  may  conclude,  therefore,  that  cones  are  even 
more  important  to  vision  than  rods.  3.  If  a  small  lighted  candle  be  moved 
to  and  fro  at  the  side  of  and  close  to  one  eye  in  a  dark  room  while  the  eyes 
look  steadily  forward  into  the  darkness,  a  remarkable  branching  figure 
Purkinje's  figures,  is  seen  floating  before  the  eye,  consisting  of  dark  lines  on 
a  reddish  ground.  As  the  candle  moves,  the  figure  moves  in  the  opposite 
direction,  and  from  its  whole  appearance  there  can  be  no  doubt  that  it  is  a 
reversed  picture  of  the  retinal  vessels  projected  before  the  eye.  The  two 
large  branching  arteries  passing  up  and  down  from  the  optic  disc  are  clearly 
visible,  together  with  their  minutest  branches.  A  little  to  one  side  of  the 
disc  there  is  an  area  free  from  vessels.  This  corresponds  to  the  yellow 
spot,  or  macula  lutea,  figure  452.  This  remarkable  appearance  is  due  to 
shadows  of  the  retinal  vessels  cast  by  the  candle.  The  branches  of  these 
vessels  are  chiefly  distributed  in  the  nerve  fiber  and  ganglionic  layers; 
and  since  the  light  of  the  candle  falls  on  the  retinal  vessels  from  in  front, 
the  shadow  is  cast  behind  them.  It  follows  that  those  elements  of  the 
retina  which  perceive  the  shadows  must  also  lie  behind  the  vessels.  Here, 
then,  we  have  a  clear  proof  that  the  light-perceiving  elements  of  the  retina 
are  not  the  fibers  of  the  optic  nerve  forming  the  innermost  layer  of  the 
retina,  but  the  external  layers  of  the  retina,  the  rods  and  cones. 

When  light  falls  on  the  rods  and  cones  it  produces  changes  which  de- 
velop nerve  impulses  that  are  transmitted  by  the  chain  of  neurones  ex- 
tending through  the  retina,  the  optic  nerve  and  chiasma,  the  geniculate 
bodies,  etc.,  to  the  cerebral  cortex  of  the  occipital  lobe,  which  is  the  sen- 
sorium  for  visual  sensations,  figures  405  and  406.  We  have  already  seen 
that  the  eye  possesses  a  wonderful  mechanical  perfection  for  receiving  and 
focusing  light  on  definite  parts  of  the  retina.  A  comparison  of  visual 
sensations  shows  that  there  are  corresponding  qualities  in  the  sensation, 
as,  for  example,  its  intensity,  duration,  localization,  complexity,  etc. 

Duration  of  Visual  Sensations.  —  The  duration  of  the  sensation  pro- 
duced by  a  luminous  impression  on  the  retina  is  always  greater  than  that 
of  the  stimulus  which  produces  it.  However  brief  the  luminous  impression, 


INTENSITY    OF   VISUAL    SENSATIONS  741 

the  effect  on  the  retina  always  lasts  for  about  one-twentieth  of  a  second. 
Thus,  suppose  an  object  in  motion,  say  a  horse,  to  be  revealed  on  a  dark 
night  by  a  flash  of  lightning,  the  image  remaining  on  the  retina  during  the 
time  of  the  flash.  The  object  is  really  revealed  for  such  an  extremely  short 
period  (a  flash  of  lightning  being  almost  instantaneous)  that  no  appreciable 
movement  could  have  taken  place  in  the  period  during  which  the  stimulus 
was  produced  on  the  retina  of  the  observer.  The  horse  would  appear  stand- 
ing in  the  position  of  motion  for  about  a  twentieth  of  a  second,  though  he 
would  not  be  seen  to  make  any  motions.  And  the  same  fact  is  proved  in  a 
reverse  way.  The  spokes  of  a  rapidly  revolving  wheel  are  not  seen  as  dis- 
tinct objects,  because  at  every  point  of  the  field  of  vision  over  which  the  re- 
volving spokes  pass,  a  given  impression  has  not  faded  before  another  comes  to 
replace  it.  Thus  every  part  of  the  interior  of  the  wheel  appears  filled. 

The  duration  of  the  after-sensation  produced  by  an  object  is  greater  in  a 
ratio  proportionate  to  the  duration  of  the  impression  which  caused  it.  Hence, 
the  image  of  a  bright  object,  as  of  the  light  of  a  window,  may  be  perceived 
in  the  retina  for  a  brief  period,  the  positive  after-image.  If,  however,  the  pri- 
mary stimulation  is  sharp  and  intense  there  will  follow  presently  an  appear- 
ance of  the  window  in  which  all  the  contrasted  lights  are  reversed,  the 
negative  after-image. 

Intensity  of  Visual  Sensations. — It  is  quite  evident  that  the  more 
luminous  a  body  the  more  intense  is  the  stimulus  it  produces.  But  the  in- 
tensity of  the  sensation  is  not  directly  proportional  to  the  intensity  of  the 
luminosity  of  the  object.  It  is  necessary  for  light  to  have  a  certain  intensity 
before  it  can  excite  the  retina,  but  it  is  impossible  to  fix  an  arbitrary  limit 
of  the  power  of  excitability.  As  in  other  sensations  so  also  in  visual  sensa- 
tions, a  stimulus  may  be  too  feeble  to  produce  a  sensation.  If  it  be  increased 
in  amount  sufficiently,  it  reaches  a  point  that  is  intense  enough  to  produce  an 
effect;  this  is  a  minimal  or  threshold  stimulus.  The  amount  of  increase  in 
the  stimulus  that  produces  a  perceptible  change  in  the  sensation  is  at  first 
very  slight,  but  later  quite  great.  It  does  not  depend  on  the  absolute  change 
of  intensity  of  the  stimulus,  but  is  proportional  to  the  intensity  of  the  stimulus 
already  acting,  Weber's  law. 

This  law,  which  is  true  only  within  certain  limits,  may  be  best  under- 
stood by  an  example.  When  the  retina  has  been  stimulated  by  the  light  of 
one  candle,  the  light  of  two  candles  will  produce  a  difference  in  sensation 
which  can  be  easily  and  distinctly  felt.  If,  however,  the  first  stimulus  is  that 
of  an  electric  arc-light,  the  addition  of  the  light  of  a  candle  will  make  no  dif- 
ference in  the  sensation.  So,  generally,  for  an  additional  stimulus  to  be  felt, 
it  may  be  proportionately  small  if  the  original  stimulus  is  small,  and  must 
be  greater  if  the  original  stimulus  is  great.  The  stimulus  increases  as  the 
numbers  expressing  its  strength,  while  the  sensation  increases  as  the 
logarithms. 


742 


THE    SENSES 


Retinoscopy.  Everyone  is  familiar  with  the  fact  that  it  is  quite  impos- 
sible to  see  the  fundus  or  back  of  another  person's  eye  by  simply  looking 
into  it.  The  interior  of  the  eye  forms  a  perfectly  black  background  to  the 
pupil.  The  same  remark  applies  to  an  ordinary  photographic  camera, 

A 

a,  b 


FIG.  468. — Diagram  to  Illustrate  the  Action  of  the  Ophthalmoscope  when  a  Plane 
Concave  Glass  is  Used,  c,  Observer's  eye.  The  light  reflected  from  any  point,  d,  on 
retina  of  a,  would  naturally  be  focused  at  e;  if  the  lens  b  is  used  is  would  be  focused  at  **, 
in  other  words,  at  back  of  c.  The  image  would  be  enlarged,  as  though  of  g,  and  would  be 
inverted.  (After  McGregor  Robertson.) 

and  may  be  illustrated  by  the  difficulty  we  experience  in  seeing  into  a  room 
from  the  street  through  the  window  unless  the  room  be  lighted  from 
within.  In  the  case  of  the  eye  this  fact  is  partly  due  to  the  feebleness  of 
the  light  reflected  from  the  retina,  most  of  it  being  absorbed  by  the  retinal 
pigment.  But  the  difficulty  is  due  more  to  the  fact  that  every  such  ray  is 
reflected  back  to  the  source  of  light  and  cannot  be  seen  by  the  unaided  eye 
without  intercepting  the  incident  light  as  well  as  the  reflected  rays  from 
the  retina.  The  difficulty  is  surmounted  by  the  use  of  the  ophthalmoscope. 

The  ophthalmoscope,  brought  into  use  by  Helmholtz,  consists  in  its  sim- 
plest form  of  a  concave  mirror  with  a  hole  in  it.  The  one  described  is  one 
of  the  less  intricate  of  the  modern  instruments.  It  consists  of,  a,  a  slightly 
concave  mirror  of  metal  or  silvered  glass  perforated  in  the  center,  and  fixed 


a 


FIG.  469. — Diagram  to  Illustrate  Action  of  Ophthalmoscope  when  a  Biconvex  Glass  is 
Used.  The  figure  d  on  retina  of  a  is  under  ordinary  conditions  focused  at  /  and  inverted. 
If  the  lens  b  be  placed  between  eyes,  the  image  h  is  seen  by  the  eye  c  as  an  enlarged  image. 
(After  McGregor  Robertson.) 

into  a  handle ;  and  b,  a  biconvex  lens  of  6  to  8  cm.  focal  length.  Two  methods 
of  examining  the  eye  with  this  instrument  are  in  common  use — the  direct 
and  the  indirect:  both  methods  of  investigation  should  be  employed.  A 
normal  eye  should  be  examined.  A  drop  of  a  solution  of  atropine  (two  grains 
to  the  ounce)  or  of  homatropine  hydrobromate  should  be  dropped  into  the  right 
eye  only  about  twenty  minutes  before  the  examination  is  commenced;  the 
ciliary  muscle  is  thereby  paralyzed,  the  power  of  accommodation  is  abolished, 


THE    OPTHALMOSCOPE 


743 


and  the  pupil  is  dilated.  This  will  materially  facilitate  the  examination; 
but  it  is  quite  possible  to  observe  all  the  details  to  be  presently  described  with- 
out the  use  of  this  drug.  The  room  being  now  darkened,  the  observer  seats 
himself  in  front  of  the  person  whose  eye  he  is  about  to  examine,  placing  himself 
upon  a  somewhat  higher  level.  A  subdued  but  steady  light  is  placed  close  to 
the  left  ear  of  the  patient  in  the  examination  of  the  right  eye.  Guiding  the 
mirror  in  his  right  hand,  and  looking  through  the  central  hole,  the  operator 
directs  a  beam  of  light  into  the  eye  of  the  patient. 
A  red  glare,  called  in  practice  the  reflex,  due  to  the 
illumination  of  the  retina,  is  seen.  The  patient  is 
then  told  to  look  at  the  little  finger  of  the  observer's 
right  hand  as  he  holds  the  mirror;  to  effect  this  the 
eye  is  rotated  somewhat  inward,  and  at  the  same 
time  the  reflex  changes  from  red  to  a  lighter  color, 
owing  to  the  reflection  from  the  optic  disc.  The 
observer  now  approximates  the  mirror,  and  with  it 
his  eye  to  the  eye  of  the  patient,  taking  care  to  keep 
the  light  fixed  upon  the  pupil,  so  as  not  to  lose  the 
reflex.  At  a  certain  distance,  which  varies  with 
the  refractive  power  in  different  eyes,  but  is  usually 
an  interval  of  about  two  or  three  inches  between 
the  observed  and  the  observing  eye,  the  vessels  of 
the  retina  will  become  visible  as  lines  running  in 
different  directions.  The  smaller  and  brighter  red 
arteries  can  be  distinguished  from  the  larger  and 
darker  colored  veins.  An  examination  of  the 
fundus  of  the  eye  reveals  the  optic  disc  and  the 
entrance  of  the  blood  vessels,  the  macula  lutea, 
and  the  fovea  centralis.  No  blood  vessels  are  seen 
in  the  fovea.  This  constitutes  the  direct  method 
of  examination,  figure  468;  by  it  the  various  details 
of  the  fundus  are  seen  as  they  really  exist,  and  it  is 
this  method  which  should  be  adopted  for  ordinary 
use. 

If  the  observer  is  ametropic,  i.e.,  is  myopic  or 
hypermetropic,  he    will  be  unable  to  employ  the 

direct  method  of  examination  until  he  has  remedied  his  defective  vision  by 
the  use  of  proper  glasses. 

In  the  indirect  method,  figure  469,  the  patient  is  placed  as  before,  and  the 
operator  holds  the  mirror  in  his  right  hand  at  a  distance  of  30  to  40  cm.  from 
the  patient's  right  eye.  At  the  same  time  he  rests  his  left  little  finger  lightly 
upon  the  patient's  right  temple,  and  holding  the  lens  between  his  thumb  and 
forefinger,  two  or  three  inches  in  front  of  the  patient's  eye,  directs  the  light 
through  the  lens  into  the  eye.  The  red  reflex,  and  subsequently  the  white  one, 
having  been  gained,  the  operator  slowly  moves  his  mirror,  and  with  it  his  eye, 
toward  or  away  from  the  face  of  the  patient,  until  the  outline  of  one  of  the 
retinal  vessels  becomes  visible,  when  very  slight  movements  on  the  part  of  the 
operator  will  suffice  to  bring  into  view  the  details  of  the  fundus  above  described, 
but  the  image  will  be  much  smaller  and  inverted.  The  lens  should  be  kept 
at  a  fixed  distance  of  two  or  three  inches,  the  mirror  being  alone  moved  until 
the  disc  becomes  visible:  should  the  image  of  the  mirror  obscure  the  disc, 
the  lens  may  be  slightly  tilted. 


FIG.  470.— The  Ophthal- 
moscope. The  small  upper 
mirror  is  for  direct,  the 
larger  for  indirect,  illumi- 
nation. 


744 


THE    SENSES 


The  Field  of  Vision. — The  field  of  vision  of  an  eye  is  that  part  of  the 
external  world  which  can  be  seen  by  it  when  the  eye  is  fixed.  Under  such 
circumstances  objects  near  the  axis  of  vision  stimulate  points  in  the  retina 
near  the  fovea  or  on  it,  while  objects  at  an  angle  of  60°  to  90°  from  the  axis 
of  vision  stimulate  regions  of  the  opposite  side  of  the  retinal  cup,  i.e.,  the 
retinal  field  is  inverted. 

The  perimeter  is  an  instrument  for  measuring  the  field  of  vision  in  terms 
of  angular  measure.  When  a  field  is  charted  by  means  of  the  perimeter  it 
is  revealed  that  objects  can  be  seen  further  out  in  the  field  in  some  directions 


330 ' 


270  a 


FIG.  471. — Perimeter  Chart,  Showing  Extent  of  Field  of  Vision  for  White  Light  and  to  the 
Colors  Red,  Green,  Yellow,  and  Blue.     (Krapart.) 

than  in  others.  For  example,  objects  in  the  temporal  field  can  be  seen  at 
an  angle  of  90°  to  100°,  while  on  the  nasal  side  they  are  seen  only  60°  to  70° 
If  the  head  is  turned  to  the  right  or  the  left  while  keeping  the  eye  fixed,  it  is 
found  that  objects  are  seen  at  a  greater  angle.  This  shows  that  the  limita- 
tions are  due  to  the  facial  boundaries  of  the  eye  preventing  the  light  from 
entering  the  eye  and  not  from  lack  of  sensitiveness  of  the  retina.  In  fact, 
the  retina  is  sensitive  to  light  out  to  the  ora  serrata. 

Localization  in  the  Retina. — Careful  exploration  of  the  retina  with 
the  perimeter  gives  a  measure  not  only  of  the  extent  of  the  visual  field,  but  of 
its  acuteness  and  localization  in  different  areas  toward  the  periphery.  Con- 
sidering the  minimal  distance  apart  which  two  luminous  points  must  be  to 
be  distinguished  as  two,  it  is  found  that  when  the  image  falls  on  the  fovea 


VISUAL   PURPLE  745 

the  two  points  may  be  very  near  together,  as  little  as  one  minute  or  even  less. 
Two  stars  can  be  seen  only  at  a  somewhat  greater  angular  distance,  two  to- 
three  minutes.  One  minute  angular  measure  covers  an  area  on  the  retina  of 
a  trifle  over  4/4.  The  diameter  of  the  cones  is  about  2/y.,  so  that  the  stimuli  in 
the  fovea  fall  on  at  least  two  separate  cones.  The  inference  seems  reason- 
able that  the  retina  in  its  most  sensitive  part  can  localize  stimuli  that  fall  on 
adjacent  cones. 

The  area  of  the  fovea  centralis  is  small,  from  o .  5  to  i .  5  mm.  Outside  of 
its  area  the  acuteness  of  vision  quickly  falls  off.  The  fact  is  roughly  esti- 
mated by  fixing  the  vision  on  a  letter  in  the  printed  line  in  the  book  before  the 
reader  and  then  determining  the  number  of  letters  to  either  side  that  can  be 
identified.  The  height  of  these  letters  is  1.5  mm.;  by  measuring  the  dis- 
tance of  the  page  from  the  eye  one  can  quickly  calculate  the  area  of  distinct 
vision  on  the  retina.  Test  types  are  printed  on  the  basis  of  an  angle  of  five 
minutes. 

In  the  outer  limits  of  the  retina  the  power  of  localizing  stimuli  is  very 
slight;  in  fact,  in  the  extreme  borders  of  the  field  it  is  difficult  to  determine 
other  than  general  form. 

Visual  Purple. — The  method  by  which  a  ray  of  light  is  able  to  stimulate 
the  endings  of  the  optic  nerve  in  the  retina  is  not  yet  understood.  It  is  sup- 
posed that  the  change  effected  by  the  agency  of  the  light  which  falls  upon  the 
retina  is  in  fact  a  chemical  alteration  in  the  protoplasm,  and  that  this  change 
initiates  a  nerve  impulse  that  is  transferred  to  the  optic  nerve  endings.  The 
discovery  of  a  certain  temporary  reddish-purple  pigmentation  of  the  outer 
limbs  of  the  retinal  rods  in  certain  animals,  e.g.,  frogs,  which  had  been 
killed  in  the  dark,  forming  the  so-called  rhodopsin  or  visual  purple,  appeared 
likely  to  offer  some  explanation  of  the  matter,  especially  as  it  was  also  found 
that  the  pigmentation  disappeared  when  the  retina  was  exposed  to  light  and 
reappeared  when  the  light  was  removed,  and  that  it  underwent  distinct 
changes  of  color  when  other  than  white  light  was  used.  It  was  also  found 
that  if  the  operation  were  performed  quickly  enough  and  in  the  dark,  the 
image  of  an  object,  optogram,  might  be  fixed  in  the  pigment  on  the  retina  by 
soaking  the  retina  of  an  animal  in  alum  solution. 

The  visual  purple  cannot,  however,  be  absolutely  essential  to  the  pro- 
duction of  visual  sensations,  as  it  is  absent  from  the  retinal  cones,  and  from 
the  macula  lutea  and  fovea  centralis  of  the  human  retina,  and  does  not  appear 
to  exist  at  all  in  the  retinae  of  some  animals,  e.g.,  bat,  dove,  and  hen,  which 
are,  nevertheless,  possessed  of  good  vision. 

However,  the  fact  remains  that  light  falling  upon  the  retina  bleaches  the 
visual  purple,  and  this  must  be  considered  as  one  of  its  effects.  It  has  been 
found  that  certain  pigments,  also  sensitive  to  light,  are  contained  in  the  inner 
segments  of  the  cones.  These  colored  bodies  are  said  to  be  oil  globules  of 
various  colors — red,  green,  and  yellow — called  chromophanes,  and  are  found 


746 


THE    SENSES 


only  in  the  retinae  of  animals  other  than  mammals.  The  rhodopsin  at  any 
rate  appears  to  be  derived  in  some  way  from  the  retinal  pigment,  since  the 
color  is  not  renewed  after  bleaching  if  the  retina  be  detached  from  its  pig- 
ment layer.  The  second  change  produced  by  the  action  of  light  upon  the 
retina  is  the  movement  of  the  pigment  cells.  On  the  stimulation  by  light 
the  granules  of  pigment  in  the  cells  which  overlie  the  outer  part  of  the  rod 
and  cone  layer  of  the  retina  become  diffused  into  the  parts  of  the  cells  be- 
tween the  rods  and  cones,  the  melanin  granules,  as  they  are  called,  passing 
down  into  the  processes  of  the  pigment  cells  A  movement  of  the  cones  and 


FIG.  472. — Sections  of  Frog's  Retina  Showing  the  Action  of  Light  upon  the  Pigment 
Cells  and  upon  the  Rods  and  Cones,  (von  Gendesen-Stort.)  A,  From  a  frog  which  had 
been  kept  in  the  dark  for  some  hours  before  death;  B,  from  a  frog  which  had  been  exposed 
to  light  just  before  being  killed.  Three  pigment  cells  are  shown  in  each  section.  In  A 
the  pigment  is  collected  toward  the  nucleated  part  of  the  cell,  in  B  it  extends  nearly  to  the 
basis  of  the  rods.  In  A  the  rods,  outer  segments,  were  colored  red  (the  detached  one 
green);  in  B  they  had  become  bleached.  In  A  the  cones,  which  in  the  frog  are  much 
smaller  than  the  rods,  are  mostly  elongated;  in  B  they  are  all  contracted. 

possibly  of  the  rods  is  also  said  to  occur,  as  has  been  already  incidentally 
mentioned.  Under  the  influence  of  the  stimulus  of  light  the  outer  parts  of 
the  cones,  which  in  an  eye  protected  from  light  extend  to  the  pigment  layer, 
are  retracted.  In  is  even  thought  by  some  that  the  contraction  is  under 
the  control  of  the  nervous  system.  Finally,  according  to  the  careful  re- 
searches of  Dewar  and  McKendrick,  and  of  Holmgren,  it  appears  that  the 
stimulus  of  light  is  able  to  produce  an  action  current  in  the  retina.  Mc- 
Kendrick believes  that  this  is  the  electrical,  expression  of  those  chemical 
changes  in  the  retina  of  which  we  have  already  spoken. 

Color  Sensations. — When  a  ray  of  sunlight  enters  the  eye  it  produces  a 
sensation  of  white  light.  But  if  the  ray  first  passes  through  a  prism,  then  it 


COMPLEMENTAL    COLORS,    AND    AFTER-IMAGES    OF    COLOR        747 

produces  sensations  corresponding  to  the  colors  of  the  spectrum.  As  is  well 
known,  white  light  is  produced  by  vibrations  of  the  luminiferous  ether  through 
a  wide  range  of  vibration  rates.  When  a  beam  of  white  light  is  passed 
through  a  dispersing .  prism  those  vibration  rates  of  low  frequency  are  re- 
fracted less  than  those  of  higher  frequency,  giving  rise  to  the  spectrum. 
Vibrations  of  the  luminiferous  ether  of  rates  just  outside  of  the  spectral  rates 
exist,  those  which  have  a  lower  rate  giving  rise  to  heat  rays,  and  those  of 
higher  rate  to  the  so-called  actinic  or  chemical  rays,  because  they  exert  a 
powerful  chemical  action.  Those  spectral  colors  which  stimulate  the  retina 
to  produce  sensations  of  color  presumably  affect  the  retinal  elements  through 
chemical  changes  which  they  produce  there.  But  this  matter  will  be 
discussed  under  theories  of  color  vision. 

The  examination  of  color  sensations  reveals  certain  correspondences  be- 
tween the  physical  color  of  the  stimulus  and  the  resulting  color  perception. 
If  a  pure  spectral  color  be  allowed  to  fall  on  the  retina,  a  corresponding  simple 
sensation  is  produced.  If  two  colors  fall  on  the  same  portion  of  the  retina 
at  the  same  time,  a  sensation  is  produced  that  is  different  from  that  which 
occurs  when  either  color  alone  stimulates.  The  same  fact  holds  true  for 
three  colors  or  more.  In  fact,  three  spectral  colors  can  be  selected  which 
by  proper  combination  can  be  used  to  produce  sensations  of  all  the  colors  of 
the  spectrum.  Such  colors  are  called  the  fundamental  colors,  and  while 
the  choice  is  more  or  less  arbitrary,  red,  green,  and  violet  are  the  colors 
usually  considered. 

Extent  of  the  Visual  Field  for  Color. — The  retina  is  most  sensitive 
to  color  in  the  region  of  the  macula  lutea.  If  by  means  of  the  perimeter  one 
explores  the  retina  to  spectral  red,  for  example,  it  is  found  that  the  color  can 
be  identified  only  at  a  distance  of  from  30°  to  50°  from  the  macula;  the 
limits  extending  out  somewhat  farther  on  the  nasal  side  of  the  retina;  that 
is,  the  part  corresponding  to  the  temporal  visual  field.  In  the  same  way  yel- 
low can  be  identified  for  from  40°  to  70°,  blue  from  40°  to  50°.  The  visual 
field  for  green  is  quite  restricted,  usually  extending  only  from  20°  to  30°. 
The  extent  of  the  color  visual  field  varies  greatly  in  different  individuals. 

Complemental  Colors,  and  After-images  of  Color. — Certain  colors, 
when  allowed  to  stimulate  the  retina  at  the  same  time,  tend  to  neutralize 
each  other.  That  is,  they  produce  sensations  approaching  white,  usually 
some  shade  of  gray,  which  will  have  a  tinge  of  one  or  the  other  primary  colors 
according  to  the  proportion  of  stimulation.  These  pairs  of  colors  are  called 
complemental  colors.  Each  spectral  color  has  its  complemental  color,  a  fact 
that  is  represented  in  figure  473.  The  complemental  colors  of  greatest  phys- 
ical significance  are  red  and  green  (greenish-blue),  yellow  and  deep  blue 
(indigo  blue),  green  (greenish-yellow),  and  violet.  . 

Positive  after-images  of  color  exist  for  a  brief  moment,  but  the  greatest 
significance  attaches  to  the  negative  after-images.  The  negative  after-images 


74-8  THE    SENSES 

of  color  following  the  stimulus  of  colored  light  upon  the  retina  are  not  the 
sensation  of  color  produced  by  the  color  of  an  object,  but  are  the  opposite 
or  complemental  color.  The  after-image  of  red  is,  therefore,  green,  and 
that  of  green,  red;  that  of  violet,  yellow,  and  of  yellow,  violet,  etc.  The 
same  relation  holds  with  the  other  colors.  A  condition  for  the  development 
of  a  strong  after-image  is  that  the  primary  image  shall  have  continued  to  a 
certain  degree  of  fatigue.  The  colors  which  reciprocally  excite  each  other 
in  the  retina  are  those  placed  at  opposite  points  in  the  color  table,  figure  473. 
The  after-images  of  color  are  most  intense  in  the  axis  of  the  visual  field  and 
are  not  always  present  in  the  periphery  of  the  retina,  as  can  readily  be  seen 
by  examining  the  chart,  figure  471. 

Color  sensations  may  also  be  produced  by  contrast.  Thus,  a  very  small 
dull  gray  strip  of  paper,  lying  upon  an  extensive  surface  of  any  bright  color, 
does  not  appear  gray,  but  has  a  faint  tint  of  the  color  which  is  the  comple- 


Green 


(OrceniiMlw) 
Cyan-blue 


Ontnpe 


Red 


FIG.  473. — Geometrical  Color  Table  for  Determining  the  Complemental  Colors. 

ment  of  that  of  the  surrounding  surface.  A  strip  of  gray  paper  upon  a  green 
field,  for  example,  appears  to  have  a  tint  of  red,  and  when  lying  upon  a  red 
surface,  a  greenish  tint;  it  has  an  orange-colored  tint  upon  a  bright  blue 
surface,  and  a  bluish  tint  upon  an  orange-colored  surface;  yellowish  color 
upon  a  bright  violet,  and  a  violet  tint  upon  a  bright  yellow  surface.  The 
color  excited  thus  must  arise  as  an  opposite  or  antagonistic  condition  of  the 
retina,  and  the  opposite  conditions  of  which  it  thus  becomes  the  subject, 
would  seem  to  balance  each  other  by  their  reciprocal  reaction.  A  necessary 
condition  for  the  production  of  the  contrast  colors  is  that  the  part  of  the 
retina  in  which  the  new  color  is  to  be  excited  shall  be  in  a  state  of  compara- 
tive repose;  hence  the  small  object  itself  must  be  gray.  A  second  condition 
is  that  the  color  of  the  surrounding  surface  shall  be  very  bright. 

Color-blindness. — Many  persons  are  unable  to  distinguish  one  or 
mo-e  of  the  fundamental  colors,  and  therefore  have  different  perceptions 
of  the  color  combinations  from  that  of  the  normal  individual.  It  is  said  that 


THEORIES    OF    COLOR  VISION  749 

from  4  to  5  per  cent,  of  men  and  about  i  per  cent,  of  women  are  defective  in 
color  vision.  The  defect  is  called  color-blindness. 

In  very  rare  cases  complete  color-blindness  exists.  Such  individuals 
distinguish  lights  and  shades  only,  that  is,  form.  A  more  common  defect, 
however,  is  the  absence  of  one  or  more  of  the  fundamental  color  sensations, 
the  most  common  of  all  being  the  green  blind,  or  the  red-green  blind.  The 
red-green  blind  individual  cannot  distinguish  red  and  green  colored  yarns 
from  each  other  or  from  shades  of  gray  which  reflect  light  with  the  same 
intensity.  When  they  are  given  the  color  test  by  the  Holmgren  yarns,  they 
indiscriminately  mix  the  reds,  greens,  and  grays.  Cases  have  been  described 
in  which  the  individual  was  red-blind  alone,  or  green-blind  alone.  A  less 
common  color  defect  is  the  inability  to  distinguish  yellows  and  blues,  yellow- 
blue  blindness. 

Color-blindness  may  occasionally  arise  from  disease  or  accident,  but  it 
is  usually  congenital.  The  individual  often  does  not  discover  his  defect  until 
examined  especially  for  his  color  vision.  He  may  have  learned  to  apply 
the  terms  green  and  red  to  surrounding  objects,  such  as  the  grass,  bricks,  etc., 
but  he  distinguishes  these  objects  by  slight  differences  in  intensity  of  lumi- 
nation,  form,  etc.,  and  not  by  the  sensations  of  color  which  the  normal 
individual  experiences. 

Theories  of  Color  Vision. — We  have  no  way  of  determining  the 
method  by  which  the  colors  stimulate  the  retina  other  than  our  inferences 
from  indirect  evidence.  It  is  probable  that  the  energy  of  light  vibration 
is  transformed  in  the  retinal  structures  into  either  physical  or  chemical 
change,  perhaps  the  latter.  Those  interested  in  the  phenomena  of  color 
vision  generally  accept  one  of  two  theories,  or  their  modifications,  in  ex- 
planation of  the  facts. 

The  Young- Helmholtz  Theory  of  Color  Vision. — This  theory  assumes 
that  there  are  three  fundamental  sensory  elements  in  the  retina  which  cor- 
respond to  and  are  stimulated  primarily  by  the  three  primary  colors — red, 
green,  and  violet.  The  theory  in  its  present  form  further  assumes  that  each 
color-perceiving  element  is  slightly  stimulated  by  others  of  the  spectral  rays, 
as  shown  in  figure  474.  When  red  rays  fall  upon  the  retina,  they  stimulate 
the  red-perceiving  elements  strongly  and  the  green  and  violet  very  feebly. 
The  resulting  sensation  is  that  of  red.  So  also  is  it  with  green  and  violet  rays. 
When  the  retina  is  stimulated  by  both  red  and  green  rays,  the  two  correspond- 
ing color-perceiving  elements  are  strongly  stimulated.  The  resulting  color 
perception,  however,  is  a  combination  of  the  two  sensations  and  corresponds 
to  some  region  of  the  spectrum  between  the  red  and  green,  according  to  the 
relative  intensity  of  the  two  stimuli.  When  all  three  color-perceiving  ele- 
ments are  stimulated  at  the  same  time,  the  theory  assumes  that  white  light 
will  be  perceived.  In  a  similar  manner  all  the  various  color  sensations  are 
arrived  at. 


750 


THE    SENSES 


Hering's  Theory  of  Color  Vision. — This  theory  is  based  on  the  assump- 
tion that  there  are  chemical  substances  in  the  retina,  photogenic  substances, 
which  are  stimulated  by  the  colors  of  the  spectrum.  It  assumes  three  photo- 
genic substances  which  are  called  the  red-green,  the  yellow-blue,  and  the 


FIG.  474. — Diagram  to  Illustrate  the  Stimulating  Effects  of  the  Three  Primary  Colors. 
(Young-Helmholtz  theory.)  i  is  the  red;  2,  green,  and  3,  violet,  primary  color  sensations. 
The  lettering  indicates  the  colors  of  the  spectrum.  The  diagram  indicates  by  the  height 
of  the  curve  to  what  extent  the  several  primary  sensations  of  color  are  excited  by  vibrations 
of  different  wave  lengths.  (Helmholtz.) 

white-black  substances.  By  the  theory,  when  the  red-green  substance  is 
stimulated  by  red  or  green  light,  respectively,  the  former  produces  destruc- 
tive or  katabolic  changes,  the  latter  constructive  or  anabolic  changes  in  the 
substance.  When  red  light  falls  upon  the  retina,  it  produces  katabolism  in 
the  red-green  substance,  which  in  turn  develops  a  nerve  impulse  that  arouses 
rg  w  y.b 


FIG.  475. — Diagram  to  Illustrate  the  Reactions  of  the  Three  Photogenic  Substances, 
according  to  Hering's  Theory.     (Foster.) 

the  sensation  of  red.  When  green  light,  on  the  other  hand,  stimulates  the 
retina,  it  produces  anabolism  of  the  red-green  substance  and  the  sensation  of 
green.  The  same  rule  holds  with  the  other  two  substances.  It  will  be 
noticed  that  this  theory  is  based  on  the  complemental  colors. 


BINOCULAR   VISION  751 

When  we  apply  the  theories  mentioned  above  to  the  phenomena  of  color- 
contrast  and  color-blindness,  we  find  that  each  is  defective  in  some  point. 
By  the  Young-Helmholtz  theory  it  is  difficult  to  understand  the  perception 
of  the  sensation  of  black,  for  by  the  theory  black  could  be  perceived  only  as 
the  absence  of  all  colors,  and  it  is  generally  granted  that  there  is  a  distinct 
black  sensation  other  than  and  different  from  mere  darkness.  This  theory 
explains  more  satisfactorily  those  cases  of  blindness  to  one  color,  as  red- 
blindness,  for  example.  The  Hering  theory,  on  the  other  hand,  gives  us  a 
rational  explanation  for  positive  black  sensation,  and  is  particularly  appli- 
cable to  the  observed  facts  of  color-contrast  and  negative  color  after-images. 

Color  after-images,  as  for  instance  the  after-images  of  green  following 
stimulation  by  red  light,  are  readily  explained  by  Bering's  theory,  since  the 
strong  katabolism  in  the  red-green  substance  will  be  followed  immediately 
by  anabolism  to  bring  this  substance  up  to  its  normal  in  the  eye,  thus  pro- 
ducing the  after-image.  This  phenomenon  can  be  explained  by  the  Young- 
Helmholtz  theory  only  by  assuming  that  following  the  stimulation  by  red 
light  and  the  consequent  fatigue  of  red-perceiving  elements  there  is  sufficient 
light  entering  the  eye  to  stimulate  the  relatively  sensitive  green  and  violet 
perceiving  elements,  thus  producing  an  after-image.  Strong  after-images 
are  perceived  in  the  dark  room,  so  that  the  Hering  theory  is  most  applicable 
in  the  explanation  of  these  cases. 

Binocular  Vision. — When  one  looks  at  an  object  with  a  single  eye, 
the  eye  is  so  adjusted  that  the  axis  of  vision  is  directed  toward  the  object 
investigated.  This  is  called  ocular  fixation.  The  ocular  fixation  is  accom- 
plished by  the  co-ordinated  contractions  of  the  six  ocular  muscles.  Its 
purpose  is  to  bring  the  image  of  the  object  examined  in  the  external  visual 
field  as  nearly  as  possible  upon  the  macula  lutea.  In  binocular  vision  both 
eyes  are  fixed  on  the  same  point  in  the  visual  field.  A  projection  of  the 
visual  axis  of  each  eye  will  pierce  the  point  of  fixation  in  the  external  object. 
It  is  evident  that  objects  to  either  side  of  the  point  of  fixation  will  give  off 
rays  which  will  enter  the  eyes,  stimulating  fields  in  the  retina  on  the  opposite 
side  of  the  visual  axis.  An  examination  of  figure  476  will  show  that  each 
point  in  the  visual  field,  A,  B,  C,  D,  stimulates  corresponding  points, 
a,  b,  c,  d,  a',  b'y  cf,  d',  in  the  retinas  of  the  two  eyes,  a,  b,  c,  d,  and 
a'  b'j  c',  d',  are  corresponding  points  in  the  two  retinas.  When  a  and  a' 
are  stimulated  at  one  and  the  same  time,  the  resulting  sensation  is  attributed 
to  one  object  in  the  visual  field,  A,  and  these  are  corresponding  points. 
This  can  be  shown  by  pressing  one  eye  out  of  its  normal  fixation  so  that 
the  primary  axes  of  the  two  eyes  are  not  directed  toward  the  same  point. 
If  one  eye  is  pressed  lightly  by  the  thumb  while  examining  a  given  object, 
as  soon  as  the  pressure  is  applied  two  objects  will  appear.  This  phenome- 
non is  known  as  diplopia.  Diplopia  is  due  to  the  fact  that  the  images  of 
visual  objects  do  not  fall  on  corresponding  points  in  the  two  retinae. 


752 


THE    SENSES 


The  parts  of  the  retinae  in  the  two  eyes  which  thus  correspond  to  each 
other  in  the  property  of  referring  the  images  which  affect  them  simulta- 
neously to  the  same  spot  in  the  field  of  vision,  are,  in  man,  just  those  parts 
which  would  correspond  to  each  other  if  one  retina  were  placed  exactly  in 
front  of  and  over  the  other,  as  in  figure  477.  Thus,  as  we  have  noticed  in 
speaking  of  the  distribution  of  the  optic  nerve  fibers,  the  temporal  portion 
of  one  eye  corresponds  to  or  is  identical  with  the  nasal  portion  of  the  other 


FIG.  476. — Diagram  Showing  the  Symmetrical  Correspondence  of  the  Retinal  Fields. 
TV,  Nodal  point;  F,  fovea  centralis.  The  observer  is  supposed  to  be  looking  down  upon  the 
optical  apparatus  from  above.  Note  that  the  line  CD,  which  is  on  the  lower  side  of  the 
object,  is  the  upper  side  of  the  image;  and  that  the  line  BD,  which  is  the  right  side  of  the 
object,  is  the  left  side  of  the  image,  which  brings  it  at  the  inner  segment  of  the  right  retina 
and  the  outer  segment  of  the  left  retina. 

eye.  The  upper  part  of  one  retina  is  also  identical  with  the  upper  part  of 
the  other;  and  the  lower  parts  of  the  two  eyes  are  identical  with  each  other. 
The  distribution  of  the  optic  nerve  fibers  corresponds  with  the  distribution 
of  the  identical  points.  The  identical  points  on  the  upper  and  lower  parts 
of  the  retinae  may  also  be  shown  by  the  following  simple  experiment. 

Pressure  upon  any  part  of  the  ball  of  the  eye,  so  as  to  affect  the  retina, 
produces  a  luminous  circle,  seen  at  the  opposite  side  of  the  field  of  vision  to 
that  on  which  the  pressure  is  made.  If,  now,  in  a  dark  room,  we  press  with 
the  finger  at  the  upper  part  of  one  eye,  and  at  the  lower  part  of  the  other, 
two  luminous  circles  are  seen,  one  above  the  other;  so,  also,  two  figures  are 


BINOCULAR    VISION  753 

seen  when  pressure  is  made  simultaneously  on  the  outer  or  the  inner  sides 
of  both  eyes.  But  if  pressure  be  made  with  the  fingers  upon  both  eyes 
simultaneously  at  their  lower  part,  one  luminous  ring  is  seen  at  the  middle 
of  the  upper  part  of  the  field  of  vision.  If  the  pressure  be  applied  to 
the  upper  part  of  both  eyes,  a  single  luminous  circle  is  seen  in  the  middle 
of  the  field  of  vision  below.  So,  also,  if  we  press  upon  the  outer  side  of  one 
eye  and  upon  the  inner  side  of  the  other  eye,  a  single  luminous  spot  is  pro- 
duced, and  is  apparent  at  the  extreme  right  of  the  field  of  vision.  The 
hemispheres  of  the  two  retinae  may,  therefore,  be  regarded  as  lying  one  over 
the  other,  as  in  C,  figure  477.  If  the  axes  of  the  eyes,  A  and  B,  figure  478, 


FIG.  477.  FIG.  478. 

FIG.  477. — Diagram  to  Show  the  Corresponding  Parts  of  the  Retinae. 
FIG.  478. — Diagram  to  Show  the  Simultaneous  Action  of  the  Eyes  in  Viewing  Objects 
in  Different  Directions. 

be  so  directed  that  they  meet  at  a,  an  object  at  a  will  be  seen  singly,  for 
the  point  a  of  the  one  retina  and  a'  of  the  other  are  identical.  So,  also,  if 
the  object  /?  be  so  situated  that  its  image  falls  in  both  eyes  at  the  same  dis- 
tance from  the  central  point  of  the  retina. — namely,  at  b  in  the  one  eye 
and  at  bf  in  the  other — /?  will  be  seen  single,  for  it  affects  identical  parts  of 
the  two  retinae.  The  same  will  apply  to  the  object  f 

The  reason  why  the  impressions  on  the  identical  points  of  the  two  retinae 
give  rise  to  but  one  sensation,  and  the  perception  of  but  a  single  image, 
must  either  lie  in  the  structural  organization  and  relations  of  the  deeper 
or  cerebral  portions  of  the  visual  apparatus,  or  it  must  be  the  result  of  a 
mental  operation;  for  in  no  other  case  is  it  the  property  of  corresponding 
nerves  of  the  two  sides  of  the  body  to  refer  their  sensations  to  one  spot. 

Many  attempts  have  been  made  to  explain  this  remarkable  relation  be- 
tween the  eyes,  by  referring  it  to  anatomical  relation  between  the  optic  nerves. 
The  circumstance  of  the  inner  portion  of  the  fibers  of  the  two  optic  nerves 
decussating  at  the  commissure,  and  passing  to  the  eye  of  the  opposite  side, 
while  the  outer  portion  of  the  fibers  continue  their  course  to  the  eyes  of  the 

48 


754  THE  SENSES 

same  side,  so  that  the  left  side  of  both  retinae  is  formed  from  one  root  of  the 
nerves,  and  the  right  side  from  the  other  root,  naturally  led  to  an  attempt 
to  explain  the  phenomenon  by  this  distribution  of  the  fibers  of  the  nerves. 
And  this  explanation  is  favored  by  cases  in  which  the  entire  half  of  one  side 
of  the  retina  sometimes  becomes  insensible. 

Visual  Judgments. — Form  and  Solidity. — The  estimation  of  the  form 
of  bodies  by  sight  is  the  result  partly  of  the  visual  sensations  and  partly  of  the 
association  of  ideas.  The  form  of  the  image  perceived  by  the  retina  depends 
wholly  on  the  outline  of  the  part  of  the  retina  affected;  the  sensation  alone  is 
adequate  only  to  the  distinction  of  superficial  forms  from  each  other  which 
lie  in  one  plane,  as  of  a  square  from  a  circle.  But  the  idea  of  a  solid  body,  as 
a  sphere,  or  a  body  of  three  or  more  surfaces,  e.g.,  a  cube,  can  be  attained 
only  by  the  action  of  the  mind  in  constructing  it  from  the  different  superficial 
images  seen  in  different  positions  of  the  eye  with  regard  to  the  object,  and 
(as  shown  by  Wheatstone  and  illustrated  in  the  stereoscope),  from  two  dif- 
ferent perspective  projections  of  the  body  being  presented  simultaneously  to 
the  mind  by  the  two  eyes.  Hence,  when,  in  adult  age,  sight  is  suddenly 
restored  to  persons  blind  from  infancy,  all  objects  in  the  field  of  vision  appear 
at  first  as  if  painted  flat  on  one  surface;  and  no  idea  of  solidity  is  formed  until 
atter  long  exercise  of  the  sense  of  vision  combined  with  that  of  touch.  The 
clearness  with  which  an  object  is  perceived,  irrespective  of  accommodation, 
would  appear  to  depend  largely  on  the  definiteness  of  stimulation  of  the  rods 
and  cones  which  its  retinal  image  covers.  Hence,  the  nearer  an  object  is  to 
the  eye,  within  the  limits  of  vision,  the  more  clearly  are  all  its  details  seen. 
Moreover,  if  we  want  carefully  to  examine  any  object,  we  always  direct  the 
eyes  straight  toward  it,  so  that  its  image  shall  fall  on  the  yellow  spot, 
which  has  already  been  shown  to  be  the  area  of  the  most  acute  vision. 

In  binocular  vision  the  images  of  an  object,  while  they  fall  in  approxi- 
mately corresponding  points  on  the  two  retinae,  are  never  absolutely  the 
same. 

When  an  object  is  placed  so  near  the  eyes  that  to  view  it  the  optic  axes 
must  converge,  a  different  perspective  projection  of  it  is  seen  by  each  eye, 
these  perspectives  being  more  dissimilar  as  the  convergence  of  the  optic  axes 
becomes  greater.  Thus,  if  any  figure  of  three  dimensions,  an  outline  cube, 
for  example,  be  held  at  a  moderate  distance  before  the  eyes,  and  viewed  with 
each  eye  successively  while  the  head  is  kept  perfectly  steady,  A,  figure  479, 
will  be  the  picture  presented  to  the  right  eye,  and  B  that  seen  by  the  left  eye. 
Wheatstone  has  shown  that  on  this  circumstance  depends  in  a  great  measure 
our  conviction  of  the  solidity  of  an  object,  or  of  its  projection  in  relief.  If 
different  perspective  drawings  of  a  solid  body,  one  representing  the  image 
seen  by  the  right  eye,  the  other  that  seen  by  the  left,  for  example,  the  drawing 
of  a  cube,  A,  B,  figure  479,  be  presented  to  corresponding  parts  of  the  two 
retinae,  as  may  readily  be  done  by  means  of  the  stereoscope,  the  mind  will 


JUDGMENTS    OF    SIZE    AND    DISTANCE 


755 


perceive  not  merely  a  single  representation  of  the  object,  but  a  body  pro- 
jecting in  relief,  the  exact  counterpart  of  that  from  which  the  drawings  were 
made. 

Judgments  of  Size  and  Distance. — The  estimation  of  the  size  of  an  object 
and  its  distance  away  from  the  observer  is  based  in  part  upon  the  visual 
image  and  in  part  upon  judgments  due  to  past  experience.  The  elements 
are  inseparable  and  mutually  dependent.  Thus,  a  lofty  mountain  many 
miles  away  may  subtend  the  same  visual  angle  as  a  small  hill  near  at  hand. 


\ 


FIG.  479. — Diagrams  to   Illustrate  how  a  Judgment  of  a  Figure  of  Three  Dimensions  is 

Obtained. 


While  the  size  and  shape  of  the  two  images  may  be  identical,  yet  the  image 
of  the  hill  near  at  hand  is  more  distinct,  its  details  are  perceived,  and  its  out- 
lines are  sharper  than  in  the  image  of  the  mountain.  If  the  atmosphere 
be  charged  with  moisture  or  with  dust,  the  image  of  the  mountain  will  be 
still  more  indistinct  and  dim.  From  previous  experiences  we  have  learned 
that  the  dimness  and  indistinctness  of  the  one  and  the  definiteness  of  the 
other  are  associated  with  distance. 

If  two  objects  are  very  near  at  hand  then  there  will  be  a  difference  in 
the  convergence  of  the  two  eyes  in  binocular  vision.  It  is  now  well  known 
that  the  ocular  muscles  are  possessed  of  a  very  delicate  muscle  sense.  This 
muscle  sense  leaves  the  impression  which  enables  us  to  judge  that  the  one 
object  is  nearer  and  the  other  farther.  In  the  common  and  familiar  objects 
about  us  we  have  from  long  experience  and  intimate  contact  learned  their 
actual  size  and  the  character  of  the  retinal  image  formed  at  definite  but  known 
distances.  When  such  an  object  forms  an  image  of  the  common  size  and 
usual  distinctness  on  the  retina  the  judgment  as  to  its  distance  is  quickly 
made. 

In  the  case  of  unknown  objects  which  are  associated  with  known  ob- 
jects, the  judgment  of  the  size  and  distance  of  the  latter  is  used  in  forming 
a  judgment  of  the  size  and  distance  of  the  former  by  comparison.  Many 
visual  deceptions  are  based  on  these  comparisons,  a  fact  that  is  often  taken 
advantage  of  by  photographers.  It  is  also  well  known  that  people  living  in 
a  moist,  hazy  climate  are  utterly  unable  accurately  to  estimate  distances 
when  suddenly  transferred  to  a  clear  mountain  climate. 


756 


LABORATORY  DIRECTIONS  FOR  EXPERIMENTS  ON  THE 
SENSE  ORGANS. 

i.  Touch. — Use  the  small  compasses  with  rounded  tips  provided  for 
the  purpose,  and  determine  the  power  of  localization  of  the  sense  of  touch 
as  follows:  Have  the  person  observed  close  his  eyes,  then  touch  different 
parts  of  the  skin,  of  the  hand,  arm,  face,  neck,  etc.,  and  let  the  observed  one 
announce  the  exact  point  touched. 

The  localization  can  also  be  determined  by  touching  two  points  on  the 
skin  with  the  points  of  the  compasses  separated  by  varying  distances.  Ex- 
amine especially  the  skin  on  the  forearm,  on  the  back  of  the  hand,  on  the 


FIG.  480. — Aristotle's  Experiment. 

palm  of  the  hand,  the  tips  of  the  fingers,  and  at  different  points  on  the  face, 
including  the  lips  and  tip  of  the  tongue.  Touch  these  regions  of  the  skin 
with  either  one  or  with  two  points  of  the  compasses,  and  allow  the  person 
observed  to  announce  results,  drawing  your  conclusions  according  to  the 
principle  of  trial  and  error.  Make  a  table  showing  the  power  of  local  dis- 
crimination in  the  different  regions. 

2.  Aristotle's  Touch  Experiment. — Roll  the  tips  of  the  middle  and 
index  fingers  over  a  marble  and  note  that  the  sensation  from  the  two  fingers 
is  interpreted  as  that  of  a  single  object.     Now  cross  the  fingers  and  repeat 
the  experiment.     This  time  there  is  the  sensation  of  touching  two  spheres. 

3.  Temperature  Sensations. — It  is  a  common  experience   that  the 
hand  brought  in  the  neighborhood  of  a  warm  or  a  cold  object  develops  the 
sensation  of  warmth  or  cold.     Examine  a  given  small  area  of  the  back  of 
the  hand,  or  a  square  centimeter  on  the  palmar  surface  of  the  wrist.     Use 
the  thermoesthesiometer,  or  a  pencil  with  large  diameter  of  the  carbon 
warmed  in  a  sand  bath.     Certain  points  will  give  stronger  sensation  of 
heat  than  others.     Map  these  out  carefully.     Examine  the  same  area  for 
the  cold.     A  large  number  of  cold  spots  will  be  found  and  they  will  not 
coincide  with  the  warm  spots,  figure  421. 


SENSATIONS   OF    SMELL 


757 


The  stimulation  for  the  hot  and  cold  spots  does  not  depend  upon  the 
absolute  temperature,  but  on  the  relative  temperature.  Insert  the  hand  in 
water  that  feels  lukewarm.  Place  the  same  hand  in  a  cup  of  quite  warm 
water  for  a  moment,  then  reinsert  it  in  the  lukewarm  water.  This  will  now 
feel  cold. 

4.  Sensations  of  Taste. — The  distribution  of  taste  organs  in  the  tongue 
is  shown  in  figure  481.  Examine  your  own  tongue  for  organs  of  sweet, 
acid,  saline,  and  bitter,  using  solutions  of  i  to  2  per  cent,  salt,  10  per  cent, 
sugar,  i  to  2  per  cent,  sulfuric  acid,  or  5  per  cent,  acetic  acid,  and  o.i  per 
cent,  quinine. 


ft. 


FIG.  481.— Localization  of  Taste.     Bitter ;  acid ;  salt,  —  •— — ;  sweet ;  T, 

tonsils;  FC,  foramen  cecum;  CF,  circumvallate  papillae;  FP,  fungiform  papillae.     (Hall.) 

Wipe  the  tongue  dry  and  apply  the  solution  named  from  the  tip  of  a  glass 
rod.  The  best  form  of  rod  is  about  15  cm.  long  by  o.  5  cm.  in  diameter,  and 
has  one  end  drawn  out  to  a  slender  pencil-shaped  tip  and  of  a  size  which 
will  suspend  a  very  small  drop.  Too  large  a  drop  diffuses  over  too  great  an 
area  of  the  tongue.  Occasionally  small  crystals  of  sugar,  salt,  etc.,  give 
more  satisfactory  results. 

Perform  the  experiments  on  yourself  before  a  mirror  and  map  the  re- 
sults as  shown  in  figure  481. 

If  the  experiments  are  done  with  care  certain  papillae  will  be  found  which 
give  one  or  two  of  the  taste  sensations,  but  not  all. 

5.  Sensations  of  Smell. — Quantitative  experiments  on  the  sense 
of  smell  are  difficult  to  determine.  Inhale  vapor  of  ammonia  so  dilute  that 
it  can  just  be  detected.  Note  that  the  sensation  is  strongest  at  the  moment 
of  drawing  the  vapor  into  the  nostril.  Fill  the  nostrils  with  the  diluted  vapor 
and  close  the  external  opening;  the  sensation  quickly  disappears.  Keeping 


THE   SENSES 

the  nostrils  closed,  walk  into  the  open  air,  then  inhale  fresh  air.  At  the 
moment  of  the  inhalation  of  fresh  air  the  ammonia  is  again  perceptible. 
Repeat  with  bergamot,  rose  water,  etc. 

6.  The  Limits  of  the  Sense  of  Hearing. — Use  a  set  of  tuning  forks 
for  the  purpose,  and  determine  the  lowest  vibration  per  second  which  can  be 
perceived  as  sound.     Determine  the  highest  limits  in  the  same  way. 

7.  Acuteness  of  the  Sense  of  Hearing. — Listen  to  the  vibrations  of 
a  tuning  fork,  or,  better,  to  the  ticking  of  a  watch  which  is  moved  back  and 
forth  from  the  ear.     Measure  the  distance  at  which  it  can  just  be  distin- 
guished.    This  experiment  should  be  performed  with  the  person  blindfolded, 
and  extraneous  noise  should,  of  course,  be  suppressed. 

70.  Dynamic  and  Static  Functions  of  the  Labyrinth. — Maxwell  working 
with  sharks  and  Woodworth  with  pigeons  have  demonstrated  new  points 
in  the  physiology  of  the  semicircular  canals  and  of  the  utriculus  and  saccu- 
lus.  Compensating  motions  are  produced  on  stimulating  the  sense  organs 
of  the  semicircular  canal,  easily  demonstrated  in  the  shark  and  pigeon. 
Destruction  of  these  sense  organs  results  in  loss  of  co-ordination  of  types 
depending  upon  the  organs  affected.  The  semicircular  canals  must  be 
operated  in  pairs  to  induce  complete  loss  of  co-ordination  in  the  particular 
plane. 

a.  Operate  on  the  Pigeon. — Remove  the  sensory  ampullae  of  the  hori- 
zontal semicircular  canals  on  each  side.     The  operation  is  performed  under 
careful  ether  anesthesia  and  must  be  aseptic  throughout.     Quick  recovery 
occurs.     The  animal  loses  the  power  of  controlling  movements,  the  com- 
pensatory movements,  in  the  horizontal  plane.     It  will  walk  round  and 
round  in  a  circle,  or  move  the  head  without  the  usual  quick  co-ordinative 
control  of  the  skeletal  musculature. 

b.  Remove  the  right  upper  and  the  left  lower  ampullae  in  a  second 
bird.     Study  its  disturbances  of  co-ordination  as  in  the  first. 

In  the  case  of  these  pigeons,  particular  attention  must  be  given  to 
supplying  water  and  feed  for  the  first  few  days  following  the  operation.  The 
birds  will  soon  learn  to  take  care  of  themselves.  They  do  not  recover 
power  of  co-ordination  as  generally  supposed.  This  was  shown  by  the 
demonstrations  of  Dr.  Woodworth  at  the  St.  Louis  meeting  of  the  Amer- 
ican Medical  Association,  1922.  Dr.  Woodworth's  pigeons  had  been 
operated  many  months  before. 

8.  Refraction. — Light  passes  out  from  a  luminous  point  in  straight 
lines  so  long  as  the  line  of  propagation  is  in  a  medium  of  uniform  density. 
If  the  rays  pass  from  a  transparent  mediurn  of  one  density  into  a  second 
medium  of  different  density,  they  will  usually  be  turned  out  of  their  course,  or 
refracted.     If  the  rays  enter  the  second  medium  at  right  angles  to  its  surface, 
they  will  continue  in  straight  lines,  but  if  they  enter  at  any  other  angle  they 
will  be  refracted.     If  the  second  medium  is  denser  than  the  first,  the  rays  will 


NEAR   AND    FAR   LIMITS    OF  VISION  759 

be  refracted  toward  the  perpendicular;  if  it  is  less  dense,  away  from  the 
perpendicular. 

Use  a  Hall's  refraction-measuring  apparatus  (constructed  of  a  carpen- 
ter's try  square).  Adjust  it  in  a  water-pan,  and  fill  to  the  exact  level  with 
clear  water.  Clamp  a  rule  to  the  vertical  limb  of  the  apparatus  at  an 
angle  of  45  degrees  and  in  line  with  the  axial  point  of  the  instrument. 
Read  the  horizontal  scale  of  the  instrument  along  the  edge  of  the  clamped 
rule.  Remove  the  instrument  from  the  pan,  using  care  not  to  disturb 
the  adjustment  of  the  ruler,  and  construct  the  angle  of  refraction  on  co- 
ordinate paper.  Determine  the  relation  of  the  angle  of  incidence  and  of 
refraction,  and  compute  the  refractive  index  of  the  water,  the  air  having 
a  refractive  index  of  one. 

Repeat  the  determination  using  a  block  of  glass.  Draw  two  sharp 
lines  at  right  angles  across  a  white  sheet  of  paper.  Lay  a  block  of  glass 
with  polished  surfaces  on  the  paper  with  one  margin  coinciding  with  the 
horizontal  line.  Insert  one  pin  at  the  intersection  of  the  lines  but  in  con- 
tact with  the  glass.  Insert  a  second  pin  at  the  opposite  margin  of  the 
glass  and  at  an  angle  of  about  40  degrees  from  the  intersection  of  the  lines 
on  the  paper.  Now  looking  through  the  glass  determine  a  third  point 
out  on  the  paper  away  from  the  glass  and  where  the  two  pins  seem  to  be 
in  line.  Remove  the  glass  and  construct  a  diagram  showing  the  angles 
of  incidence  and  of  refraction. 

The  index  of  refraction  is  found  by  the  formula: 

sin  i 

=  Refractive  index 

sin  r 

If  equal  distances  are  laid  off  on  the  hypotenuses  then  one  may  substitute 
the  actual  measurements  of  the  vertical  legs  for  the  sins  in  the  formula 
and  calculate.  Repeat  several  times. 

9.  To  Determine  the  Refractive  Power  of  a  Convex  Lens. — Use  a 
meter  stick  which  is  provided  with  a  movable  diaphragm  or  screen,  and  a 
holder  for  a  lens.  Measure  the  focal  distance  of  lens  number  i  as  fur- 
nished from  the  optical  set.  P.ut  the  lens  in  its  holder  and  focus  the  image 
of  the  sun  or  of  an  electric  bulb  on  the  screen,  moving  the  screen  back  and 
forth  until  the  sharp  focus  is  determined.  If  the  lens  is  accurately  ground, 
the  focus  will  be  at  a  distance  of  one  meter,  which  is  the  refractive  power  of 
a  one-diopter  lens  by  definition.  In  the  same  way  determine  the  refrac- 
tive power  of  lenses  numbers  2,  3,  and  4.  If  a  source  of  light  of  a  finite 
distance  is  used,  then  correction  must  be  made  for  the  divergence  of  the 
rays  by  the  formula: 

1  +  -  -  -1- 
f        i'  "  F 

Where  f  and  f  are  the  conjugate  foci  and  F  the  principal  focal  distances  in 
meters. 


760  THE    SENSES 

• 
Construct  a  diagram  showing  the  path  of  the  light  in  the  formation  of 

the  image  in  these  cases. 

If  the  measurement  in  the  above  case  is  made  through  two  parallel  open- 
ings or  diaphragms  about  5  mm.  in  diameter  each,  and  separated  by  i  or 
2  cm.,  the  point  of  focus  can  be  more  accurately  determined  (see  Schemer's 
Experiment,  No.  14).  Construct  the  mathematical  figure  showing  the 
course  of  both  cones  of  rays  in  this  test. 

10.  Determination  of  Near  and  Far  Limits  of  Vision. — Support  a 
meter  stick  in  a  horizontal  position  at  a  comfortable  level  for  the  eye.  Mount 


FIG.  482. — Diagram  of  Experiment  to  Ascertain  the  Minimum  Distance  of  Distinct  Vision. 

a  needle  in  a  cork  and  set  it  on  the  meter  stick  about  25  cm.  in  front  of  the 
eye.  Make  two  pin-holes  in  a  card  at  a  distance  of  about  2  mm.  from  each 
other.  Hold  this  card  with  the  pin-holes  close  in  front  of  the  right  eye,  and 
bring  the  eye  up  to  the  end  of  the  meter  stick;  cover  the  other  eye.  Ob- 
serve that  when  the  needle  is  brought  nearer  and  nearer  to  the  eye,  at  a 
certain  distance  it  becomes  double.  Determine  this  distance  very  accurately. 
It  is  the  near-point  of  accommodation  for  the  right  eye.  Make  the  same 
determination  for  the  left  eye. 

Hold  the  punctured  card  in  front  of  the  right  eye,  and  move  the  needle 
(it  is  better  to  use  something  larger)  farther  and  farther  away  until  it  becomes 
again  double,  if  it  does  so.  This  is  the  far-point  of  accommodation.  In 
normal  eyes  infinity  is  the  far  limit.  In  practice  an  eye  that  has  no  far 
limit  under  twenty  feet  is  considered  normal.  This  test  should  be  made  on 
each  eye. 

Test  near  vision  with  printed  letters  the  size  of  those  on  this  page. 
Mount  a  test  paper  on  a  small  cube  and  run  it  along  the  meter  stick  as 
above.  Test  one  eye  at  a  time.  When  the  printed  letters  begin  to  lose 
their  sharp  black  outlines  the  near-point  is  reached.  To  test  the  far 
limit  use  letters  large  enough  to  be  clearly  seen  at  a  distance  of  20  feet  by 
the  normal  eye,  see  the  test  chart. 

ii.  Inverted  Image  on  the  Retina. — Dissect  off  a  segment  of  the 
sclerotic  of  a  fresh  ox  eye,  or  use  a  fresh  eye  from  an  albino  rabbit.  Make 
a  tube  of  black  paper  of  the  size  of  the  eye,  and  insert  the  eye  in  one  end, 
with  the  cornea  directed  into  the  tube.  In  the  dark  room  examine  the  image 


SPHERICAL  ABERRATION  761 

of  the  candle  flame  as  formed  on  the  retina  of  the  eye  in  the  tube.  In  a 
favorable  experiment,  a  clear  inverted  image  of  the  candle  can  be  seen  on 
the  retina  through  the  semi-transparent  membranes  of  the  eye.  The  same 
experiment  can  be  demonstrated  with  the  camera,  or  with  a  small  lens,  using 
a  ground-glass  plate  to  make  the  image  more  apparent. 

12.  Spherical  Aberration. — In  physical  optics  it  is  found  that  it  is 
difficult  to  grind  lenses  so  that  they  will  refract  equally  in  the  center  or 
optical  axis  and  in  the  periphery.     Unequal  refraction  of  these  two  regions 
is  called  spherical  aberration.     It  is  corrected  in  optics  by  diaphragms  which 
shut  out  the  light,  either  from  the  borders  of  the  lens  or  from  its  center. 
The  former  method  is  used  in  the  eye.     To  demonstrate  spherical  aberra- 
tion, look  at  an  object  two  meters  from  the  eye,  such  as  a  part  of  the  window 
moulding.     Pass  a  card  close  in  front  of  the  eye  until  the  light  enters 
only  at  the  margin   of   the  pupil,  i.e.,  the  borders  of  the  lens.     It  will 
be  found  that  the  object  is  no  longer  in  focus  and  the  outlines  are  dim 
and  diffused.     Step  nearer  and  nearer  the  window,  when  quite  close  the 
outlines  of  the  moulding  become  clear  again.     Normal  eyes  are  near- 
sighted for  the  rays  that  are  refracted  by  the  borders  of  the  lens. 

13.  Chromatic  Aberration. — Look  toward  the  borders  between  the 
sash  and  the  bright  light  of  an  open  window,  at  a  distance  of  twenty  feet  or 
more.     Use  the  right  eye  only.     Bring  a  card  across  the  pupil  approaching 
from  the  side  of  the  light  until  the  eye  is  almost  covered  with  the  card.     The 
window  sash  will  seem  to  have  a  blue- violet  fringe.     If  the  card  is  brought 
across  from  the  opposite  side,  the  sash  will  have  a  reddish-yellow  fringe. 

Make  a  cross  of  two  strips  of  Bradley' s  pure  color  paper,  one  red  and 
the  other  blue,  on  a  black  surface.  When  held  at  the  proper  distance 
the  red  appears  nearer  than  the  blue.  This  phenomenon  is  brought  out 
more  strongly  by  covering  the  colored  papers  with  very  thin  white  tissue 
paper.  The  judgment  of  distance  is  based  on  the  effort  of  accommoda- 
tion which  is  greater  for  the  red  than  for  the  blue  and  violet  rays. 

14.  Schemer's  Experiment. — Use  two  needles  on  corks,  the  method  de- 
scribed in  Experiment  10,  placing  one  at  a  distance  of  20  cm.,  and  the  other 
about  60  cm.  from  the  eye.     Use  only  the  right  eye,  look  through  two  pin- 
holes  in  a  card  at  the  far  needle.     The  near  needle  will  appear  double 
but  the  images  will  be  somewhat  blurred.     While  looking  at  the  far  needle, 
bring  a  cardboard  across  the  right  hole,  note  that  the  left  image  of  the  near 
needle   disappears,   and  vice  versa.     If  one   accommodates  for  the   near 
needle,  the  far  needle  appears  double,  and  upon  covering  the  right  hole  with 
the  card  the  right  image  of  the  far  needle  disappears.     This  is  known  as 
Scheiner's  Experiment.     It  depends  on  the  diffuse  stimulation  of  points 
on  the  retina  outside  of  the  principal  axis.     The  apparent  images  are 
referred  out  in  space  along  the  corresponding  secondary  axes.     Construct 
a  diagram  to  explain  these  phenomena. 


762  THE    SENSES 

15.  Purkinje-Sanson's  Images. — Examine  the  eye  of  another  person 
in  a  dark  room  as  follows:     With  the  observing  eye  focus  for  a  far  object, 
let  the  observer  hold  a  candle  slightly  to  one  side  of  the  axis  of  vision  and 
about  one  foot  from  the  eye.     If  the  observer  looks  into  the  other  eye  from 
the  side  opposite  the  candle,  he  will  be  able  to  see  three  reflected  images, 
figures  457  and  45 8.     One,  from  the  anterior  surface  of   the   cornea,  is 
bright  and  distinct,  and  of  medium  size  and  erect.     In  the  middle  of  the 
pupil  there  will  be  a  second  image,  larger  and  quite  dim.     This  is  a  reflec- 
tion from  the  front  of  the  lens.     The  third  image,  reflected  from  the  posterior 
surface  of  the  lens,  will  seem  to  be  farther  back  in  the  eye,  quite  small  and 
inverted.     These  images  can  all  three  be  seen  at  once  with  careful  adjust- 
ment of  the  relative  positions  of  the  candle  and  the  observer,  with  refer- 
ence to  the  axis  of  vision  of  the  eye  observed. 

If  the  observer  protects  his  own  eye  from  the  direct  light  of  the  candle 
fay  a  blackened  cardboard  between  his  eye  and  the  candle,  and  asks  the 
observed  person  to  accommodate  now  for  near  objects,  now  for  far,  keeping 
the  axis  of  vision  constant,  he  will  be  able  to  note  that  the  middle  image, 
•i.e.,  the  one  from  the  anterior  surface  of  the  lens,  changes  in  size  and  in 
relative  position  with  reference  to  the  other  two,  which  are  essentially  con- 
•stant.  With  near  accommodation  this  image  becomes  smaller  and  seems 
to  move  toward  the  image  from  the  cornea;  with  far  accommodation  it 
becomes  larger  and  appears  to  move  to  the  image  reflected  from  the  posterior 
surface  of  the  lens.  This  shows  that  the  act  of  accommodation  consists  in  a 
change  in  the  convexity  of  the  front  of  the  lens. 

16.  The  Phakoscope  of  Helmholtz. — This  classical  instrument  was 
invented  by  Helmholtz  to  demonstrate  the  act  of  accommodation,  as  out- 
lined in  the  second  paragraph  of  the  preceding  experiment.     Repeat  the 
preceding  experiment,  using  this  instrument  in  a  dark  room. 

17.  Astigmatism. — Astigmatism  is  a  term  used  to  describe  the  con- 
dition  of   unequal   curvature  of  the  refracting  surfaces  of  the  eye  in  the 
different  meridia.    The  cornea  is  the  surface  which  usually  shows  the  greatest 
astigmatism.     The  defect  is  demonstrated  by  numerous  forms  of  astigmatic 
charts,  the  most  serviceable  of  which  are  the  barred-letter  test  type,  the  clock 
dial,  or  the  dials  shown  in  figure  463  or  483.     Hang  an  astigmatic  dial  at  a 
distance  of  six  meters  and  test  the  right  and  left  eyes  separately,  as  follows: 
When  the  vision  is  focused  on  the  center  of  the  dial,  if  the  eye  is  normal, 
the  three  bars  in  each  radius  of  the  clock  dial  will  be  seen  with  equal  distinct- 
ness and  have  sharp  black  lines.     In  an  astigmatic  eye  one  or  more  of  these 
radii  will  appear  sharp  and  distinct,  while  the  other  will  appear  dim  and 
indistinct,  the  relative  difference  depending  upon  the  degree  of  astigmatism. 
Note  the  meridian  of  astigmatism  in  the  right  and  left  eyes  separately.     Use 


THE  BLIND    SPOT 


763 


the  test  set,  and  find  the  cylinder  necessary  to  correct  the  astigmatism  in 
each  eye  and  determine  its  meridian. 

Astigmatism  is  commonly  shown  by  the  presence  of  radii  when  one  looks 
at  the  stars  at  night,  or  by  the  ragged  outline  of  a  pin-hole  in  a  card,  when 


FIG.  483. — Disc  of  Concentric  Lines  for  the  Astigmatic  Test. 

held  at  arm's  length  against  a  white  sky.  In  extreme  cases  outlines  like 
the  bars  in  the  window  sash  or  checks  in  clothing  may  be  distorted,  or  some 
of  the  lines  may  not  even  be  seen. 

1 8.  The  Blind  Spot. — Look  with  the  right  eye  at  the  spot  in  the  ac- 
companying figure  at  a  distance  of  about  20  to  25  cm.,  covering  the  left  eye. 
Hold  the  spot  in  the  line  of  direct  vision  and  move  the  book  to  and  from 
the  eye;  in  some  cases  it  is  necessary  to  rotate  the  book  slightly.  It  will  be 
found  that  the  cross  to  the  right  will,  at  a  certain  position,  completely  dis- 
appear. This  happens  when  its  image  falls  on  the  retina  directly  over  the 
entrance  of  the  optic  nerve,  which  has  no  visual  cells,  and  is,  therefore,  the 
blind  spot.  This  area  is  large  enough  to  cause  a  man  completely  to 
disappear  from  the  field  of  vision  at  a  distance  of  about  one  hundred 
meters. 

Place  a  sheet  of  white  paper  at  a  distance  of  30  cm.  in  front  of  the  eye, 
holding  the  head  in  a  fixed  position  against  the  special  support  furnished; 
look  with  the  right  eye  at  the  top  of  the  cross  made  on  the  left  of  the  sheet 


FIG.  484. — Diagram  for  Demonstrating  the  Blind  Spot. 

of  paper.  Covering  the  sharpened  portion  of  a  lead  pencil  with  white 
paper,  leaving  the  black  tip  exposed,  move  this  pencil  across  the  paper 
from  the  visual  center  to  the  right.  At  a  certain  distance  the  black  lead 
will  suddenly  disappear.  Mark  this  point.  Continue  to  move  the  pencil 
until  the  lead  reappears.  Mark  this  point.  These  two  points  represent 


764 


THE    SENSES 


the  limits  of  the  blind  spot  in  the  horizontal  plane,  as  magnified  by  the 
conditions  of  the  experiment.  Mark  the  limits  in  the  other  meridians  in 
the  same  manner.  Compute  from  the  figures  obtained  the  exact  size  of 


FIG.  485.— The  Blind  Spot  with  the  Eye  30  cm.  from  the  Paper.     The  irregularity  of 
outline  is  due  to  the  larger  blood  vessels. 

the  blind  spot  in  your  right  eye,  figure  485.  Repeat  on  the  left  eye.  Usually 
these  areas  are  not  symmetrical.  The  computation  may  be  based  on  the 
following  proportion:  a,  the  distance  of  the  map  from  the  nodal  point  of  the 
eye,  is  to  b,  the  diameter  of  the  map  of  the  blind  spot,  as  c,  the  distance  from 
the  nodal  point  to  the  retina,  which  is  i.  5  cm.,  is  to  x,  the  diameter  of  the 
actual  blind  spot  in  the  retina,  a  :  b  : :  c  :  x.  x  varies  from  i .  5  to  3  or  more 
mm. 

19.  Relations  of  the  Size  of  the  Retinal  Image  to  Distance. — Com- 
pute the  size  of  the  retinal  images  of  familiar  objects  by  the  equation  given 
in  the  last  experiment.     Compute  the  size  of  the  image  formed  on  the  retina 
by  a  man  six  feet  tall  at  a  distance  of  100  feet.     Compute  the  size  of  the 
image  formed  by  a  tower  125  feet  tall  at  a  distance  of  575  feet. 

20.  Purkinje's    Shadows. — Stand    in   front  of  a  blackened  wall  in 
the  dark  room.     While  looking  toward  the  wall  with  the  right  eye  accom- 
modated for  distant  objects,  move  a  lighted  candle  back  and  forth  about 
10  to  20  cm.  to  the  right  of  the  eye  and  a  little  below  its  level.     Presently 
many  branching  shadows  will  be  seen  as  though  they  stood  in  space  in  front 
of  the  individual.     These  are  the  shadows  of  the  blood  vessels  cast  upon  the 
retina.     A  careful  examination  will  show  that  these  shadows  seem  to  con- 
verge to  a  point  to  the  right  of  the  center  of  vision  of  the  right  eye.     By 
moving  the  candle  up  and  down  or  from  side  to  side,  the  shadows  seem  also  to 
move  slightly.     Many  persons  can  readily  see  Purkinje's  figures  by  looking 
through  the  narrow  spaces  between  the  fingers  of  the  hand  moved  close  in 
front  of  the  eye,  when  the  vision  is  directed  toward  a  bright  sky.     One  can 
demonstrate  by  this  means  that  the  macula  is  free  from  blood  vessels,  since 


COLOR   BLINDNESS  765 

the  pattern  of  the  blood  vessels  around  the  borders  of  the  macula  is  very 
readily  determined.  This  is  especially  true  if  there  is  slight  retinal 
congestion. 

21.  Duration  of  the  Retinal  Image. — When  a  beam  of  light  falls 
upon  the  retina  for  an  instant  it  produces  a  stimulus  which  endures  for  a 
time  after  the  stimulus  is  removed.     This  interval  can  be  measured  by  the 
proper  mechanical  device.     Place  on  the  color  wheel  a  disc,  which  has  a 
small  segment  cut  out  at  one  point  on  the  periphery.     Put  a  printed  page 
behind  the  segment  with  the  observer  standing  in  front.     Rotate  the  segment 
faster  and  faster  until  the  printed  page  is  seen  continuously.     At  this  point 
the  visual  image  made  at  one  revolution  of  the  disc  lasts  until  the  next  im- 
pression on  the  same  spot.     The  speed  of  the  revolution  of  the  color  wheel 
can  be  measured  by  attaching  an  electric  contact  key  and  signal  magnet  to 
the  disc  wheel  and  measuring  the  rate  of  interruptions  against  the  known 
vibrations  of  a  tuning  fork.     The  same  phenomenon  may  be  determined  by 
placing  on  the  disc  two  complemental  colors  and  judging  the  speed  of 
revolution  required  for  complete  fusion. 

22.  Limits  of  the  Field  of  Vision. — The  limits  of  the  visual  field  are 
determined  by  direct  measurement  with  the  perimeter.     Set  the  person 
whose  retina  is  to  be  measured  in  a  comfortable  erect  position,  with  one  eye 
at  the  center  of  the  arc  of  the  perimeter  a  d  the  other  covered  by  an  eye- 
shade.     The  observed  eye  must  be  fixed  on  the  center  of  the  field  of  vision, 
and  care  must  be  used  to  prevent  obstruction  of  the  field.     The  examination 
is  made  with  greatest  accuracy  by  bringing  an  object  into  the  field  of  vision 
from  behind  the  person  observed.     When  the  individual  examined  first 
detects  the  presence  of  the  object,  he  announces  it  and  the  angle  is  read  off 
from  the  arc  of  the  perimeter  and  recorded  on  the  chart  for  the  purpose. 
These  readings  should  be  made  in  about  twelve  radii.     They  should  be 
made  for  each  eye. 

23.  Limits  for  the  Field  of  Vision  for  Color. — To  measure  the  limits 
of  the  field  of  vision  for  color  one  should  proceed  as  in  the  preceding  experi- 
ment, except  that  small  squares  of  colored  papers  are  brought  into  the  field 
from  the  rear.     The  retina  should  be  mapped  for  red,  green,  yellow,  and 
blue.     Use  Bradley' s  pure  color  papers.     Take  four  penholders  and  mount 
on  the  end  of  one  a  centimeter  square  of  red  paper,  on  the  others  green, 
yellow,  and  blue.     To  make  a  determination  bring  the  color  up  from  behind 
and,  as  soon  as  it  is  certainly  detected  and  announced,  remove  it  from  the 
field  of  vision.     Examine  the  eye  for  all  four  colors  at  one  sitting,  mixing 
them  indeterminately  in  the  individual  tests.     Occasionally  an  eye  will  be 
found  which  exhibits  a  well-marked  restriction  of  the  color  field,  though  the 
individual  himself  may  not  be  completely  color-blind. 

24.  Color-blindness.  —  Make    an    examination    for    color-blindness, 
using  Holmgren's  colored  yarns.     Spread  the  yarns  out  on  a  table  in  the  best 


766  THE   SENSES 

of  light.  Place  the  three  confusion  skeins  in  front  of  the  individual  to  be  ex- 
amined and  ask  him  to  match  them  quickly  from  the  skeins  on  the  table, 
paying  no  attention  to  lights  and  shades  of  the  same  color.  A  color-blind 
individual  will  confuse  colored  skeins,  most  usually  the  reds,  greens,  and 
grays. 

25.  Color  Mixing. — Use  Bradley' s  color  wheel  and  test  the  effect  of 
simultaneous  stimulation  of  the  retina  with  two  or  more  colors,  by  placing 
on  the  wheel  two  or  more  colored  discs,  rotating  the  wheel  at  a  speed 
sufficient  to  cause  complete  fusion.     The  sensation  produced  by  two  colors 
applied  simultaneously  will  be  entirely  different  from  that  produced  by 
either  alone.     Red  and  green  (or  greenish-blue),  when  mixed  in  the  proper 
proportion,  produce  a  sensation  of  gray.     The  same  effect  may  be    had 
from  yellow  and  blue,  orange  and  violet,  or  any  of  the  complementary 
colors  chosen  according  to  the  geometrical  color  table,   figure  473.     By 
mixing  three  colors,  red,  green,  and  violet,  in  the  proper  proportion  one 
can  produce  a  sensation  almost  the.  same  as  that  produced  by  white  light. 

26.  Color  After-images. — Color   after-images    can   be    demonstrated 
by  looking  continuously  at  the  center  of  one  of  the  primary  colors  of  Bradley' s 
color  charts  against  a  white  or  gray  wall  until  there  is  apparent  fatigue, 
then  suddenly  removing  the  chart.     An  after-image  of  approximately  the 
complementary  color  will  appear  in  the  course  of  a  few  seconds.     Occasion- 
ally these  images  are  very  vivid.     The  experiments  are  brilliant  if  performed 
in  the  dark  room,  using  colored  gelatin  screens  through  which  an  intense 
light  shines.     When  the  light  is  turned  off,  a  brilliant  after-image  of  the 
complementary  color  appears. 

27.  Retinoscopy. — IJse  the  ordinary  small  ophthalmoscope   and  ex- 
amine the  retina  of  the  eye  of  a  cat  or  rabbit.     Dilate  the  pupil  by  the  use  of 
atropine.     Place  the  animal  whose  eye  is  to  be  examined  on  a  support  in 
front  of  a  bright  but  uniform  light  (an  Argand  burner).     Reflect  the  light 
from  the  mirror  of  the  ophthalmoscope  through  the  pupil  into  the  retinal  cup 
of  the  animal.     Usually  the  ophthalmoscope  has  to  be  focused  for  a  cat's 
retina.     When  a  good  light  is  secured,  the  retinal  cup  will  appear  as  a  bril- 
liantly colored  disc,  with  the  branching  blood  vessels,  and  usually  with 
some  brilliant  bluish-green  pigment  in  the  lower  portions  of  the  retinal 
disc. 

After  some  practice  on  the  cat  or  rabbit,  the  student  should  examine 
the  retina  of  one  of  his  mates,  preferably  an  eye  that  has  an  unusually  wide 
pupil.  In  some  cases  a  light  dosage  of  homatropine  may  be  used  on  one  eye. 
This  will  dilate  the  pupil  and  the  examination  will  be  much  easier. 

Students  are  not  recommended  to  use  atropine  unless  under  conditions 
which  permit  the  eye  to  rest  for  two  or  three  days  following. 

28.  Corneoscopy. — Astigmatism  is  usually  a  defect  of  the  cornea. 
It  is  measured  most  accurately  by  the  Javal-Schiotz  pattern  of  ophthal- 


THE   TEST    SET  7^7 

moscope.  Seat  the  patient  erect  with  his  head  supported  in  the  head  rest 
and  eyes  level  with  the  instrument  and  one  covered.  Set  the  pointer  of 
the  telescope  at  90  and  focus  for  a  clear  image  of  the  mires  from  the  corneal 
surface  of  the  exposed  eye  raising  or  lowering  the  telescope  if  necessary. 

Turn  the  telescope  until  the  meridian  lines  of  the  adjacent  images  form 
an  unbroken  line  and  the  spurs  an  exact  cross.  This  is  the  primary  posi- 
tion and  the  radius  of  curvature  is  read  off  the  side  scale  in  millimeters 
and  tenths. 

Set  the  primary  position  indicator  on  the  left  side  wheel  at  "o,"  then 
turn  the  telescope  through  90  degrees  or  until  the  secondary  position  is 
found.  If  there  is  no  astigmatism  the  cross  will  still  be  perfect.  If  there 
is  astigmatism  then  adjust  the  right  hand  wheel  to  reform  the  cross  and 
read  the  right  scale  as  before  and  compute  the  astigmatism. 

29.  Visual  Acuity. — The  localizing  power  of  the  retina  is  measured  by 
the  angle  formed  at  the  nodal  point  by  the  rays  from  the  opposite  borders  or 
limits  of  an  object  that  can  just  be  identified.     The  standard  test  is  based  on 
the  size  of  an  object  at  a  distance  of  twenty  feet  which  will  subtend  an  angle 
of  one  minute.     The  letters  of  the  standard  test  chart  are  constructed  on  a 
total  angle  of  five  minutes,  but  the   identifying   marks  subtend  an  angle 
of  one  minute.     The  visual  acuity  of  the  eye  should  be  tested  first  for 
the  right  eye,  then  for  the  left.     Hang  a  test  chart  at  a  distance  of  twenty 
feet,  so  that  its  disc  is  well  illuminated,  and  allow  the  individual  tested 
to  read  off  the  letters  on  the  chart,  beginning  with  the  larger  ones  at  the  top. 
The  letters  on  this  chart  are  constructed  on  the  basis  of  a  visual  angle  of 
five  degrees.     When  the  letters  marked  "twenty  feet"  or  "six  meters" 
represent  the  limit  of  accurate  identification,,  the  visual  acuity  is  said  to 

be  — ,  i.e.  i,  or  normal.    If  the  line  marked  "  thirty  feet"  is  the  limit  that  can  be 
20 

read  at  the  normal  distance  of  twenty  feet  the  eye  is  subnormal  and  the  visual 

acuity  is  measured  by  the  fraction  — ,  i.e.  2/3  that  of  the  standard.     If 

3° 

20 
the  "fifteen feet"  test  can  be  read,  then  the  visual  acuity  is  — ,  or  4/3  the 

standard. 

If  the  eyes  tested  are  astigmatic,  or  have  other  optical  defects,  these 
must  first  be  corrected  before  testing  for  visual  acuity. 

30.  The  Test  Set. — The  student  is  recommended  to  close  the  experi- 
ments on  the  eye  by  fitting  glasses  for  himself  and  at  least  two  others.     He 
should  correct  for  the  defects  that  have  been  revealed  in  the  preceding  experi- 
ments, especially  for  astigmatism;  myopia,  or  hypermetropia;  and  presby- 
opia.    Of  course  each  eye  must  be  tested  and  fitted  separately. 


CHAPTER  XVI. 

THE  REPRODUCTIVE  ORGANS. 

THE  REPRODUCTIVE  ORGANS  OF  THE  MALE. 

THE  male  reproductive  organs  comprise  the  Testis,  the  Ductus  Deferens 
the  Vesicula  Seminalis,  the  Prostate,  and  the  Penis. 

The  Testis. — The  testis  consists  of  two  parts,  i,  the  testicle  proper, 
which  is  covered  by  the  tunica  albuginea  and  secretes  the  germinal  cells, 
and  2,  the  conducting  tubules,  which  compose  the  epididymis  and  ductus 
deferens. 

The  testicle  is  divided  by  connective-tissue  septa  into  lobules,  each  of 
which  is  an  aggregation  of  tubuli  seminiferi.  Each  tubule  is  limited  by  a 
membrana  propria  on  which  rests  the  germinal  epithelium. 


FIG.  486. 


FIG.  487. 


FIG.  486. — Plan  of  a  Vertical  Section  of  the  Testicle,  Showing  the  Arrangement  of  the 
Ducts.  The  true  length  and  diameter  of  the  ducts  have  been  disregarded,  a,  a,  Tubuli 
seminiferi  coiled  up  in  the  separate  lobes;  6,  tubuli  recti;  c,  rete  testis;  d,  ductuli  efferentes 
ending  in  the  coni  vasculosi;  /,  e,  g,  convoluted  canal  of  the  epididymis;  h,  vas  deferens;/, 
section  of  the  back  part  of  the  tunica  albuginea;  i,  i,  fibrous  processes  running  between  the 
lobes;  s,  mediastinum. 

FIG.  487. — Vertical  Section  through  the  Wall  of  the  Tubules  of  Epididymis.  X  700. 
(Kolliker.)  b,  Connective  tissue  and  smooth  muscle  cells;  e,  basal  layer  of  epithelial  cells; 
/,  high  columnar  cells;  p,  pigment  granules  in  columnar  cells;  c,  cuticula;  h,  cilia. 

The  male  reproductive  cells  are  all  descended  from  primitive  germ  cells, 
the  archispermiocytes,  that  become  differentiated  during  the  fetal  life. 

On  the  approach  of  sexual  maturity  the  process  of  spermatogenesis  begins. 
The  germinal  cells  multiply  rapidly,  and,  by  a  complex  series  of  mitotic 

768 


THE   DUCTUS   DEFERENS 


769 


divisions  or  stages,  form  ultimately  the  male  reproductive  cells,  or  sperm 
cells. 

The  important  stages  in  order  are:  archispermiocyte,  spermatogonia, 
primary  and  secondary  spermatocytes,  spermatids,  and  spermatozoa.  The 
spermatogonia  stage  is  the  stage  of  rapid  multiplication;  the  spermatocyte, 
that  of  maturation,  comparable  to  the  maturation  stage  of  the  ovum. 

The  spermatozoa,  or  sperm  cells,  are  the  essential  male  reproductive  cells. 
Each  mature  spermatozoon  consists  of  a  minute  oval  head,  a  middle  piece,  and 


SfC.l 


\ 

spg.r 

FIG.  488. — Later  Stages  in  Spermatogenesis  of  the  Bull,  spg.r,  Reserve  spermato- 
gonium;  spg,  spermatogonium;  spc.g,  spermatocyte  in  late  synapsis  stage;  spc.i,  spermacyte 
in  stage  just  preceding  the  maturation  divisions;  spd,  spermatids  in  advanced  stage  of 
histogenesis,  with  heads  deeply  embedded  in  Sertoli  cell.  Highly  magnified.  (After 
Schoenfeld.) 

a  tail.  The  head  consists  almost  entirely  of  the  cell  nucleus,  while  the  mid- 
dle piece  and  tail  are  cytoplasmic  structures.  The  head  is  4/z  by  2.5^. 
The  middle  piece  and  tail  are  about  50  to  6o/z  long.  The  tail  is  essentially 
a  cilium,  and  exhibits  the  power  of  flagellate  movement. 

The  Ductus  Deferens. — This  is  the  single  duct  proceeding  from  each 
testicle  to  join  its  fellow  at  the  base  of  the  bladder.  Each  has  an  ampulla  or 

49 


THE   REPRODUCTIVE    ORGANS 

a 


FIG.  489.— Section  of  a  Tubule  of  the  Testicle  of  a  Rat,  to  Show  the  Formation  of  the 
Spermatozoa,  a,  Spermatozoa;  b,  seminal  cells;  c,  spermatoblasts,  to  which  the  sper- 
matozoa are  still  adherent;  d,  membrana  propria;  e,  fibro-plastic  elements  of  the  connective 
tissue.  (Cadiat.) 


FIG.  490. — Dissection  of  the  Base  of  the  Bladder  and  Prostate  Gland,  Showing  the 
Vesiculae  Seminales  and  Ductus  Deferens.  a,  Lower  surface  of  the  bladder  at  the  place 
of  reflection  of  the  peritoneum;  b,  the  part  above  covered  by  the  peritoneum;  »,  left  ductus 
deferens,  ending  in  e,  the  ejaculatory  duct;  the  ductus  deferens  has  been  divided  near  i, 
and  all  except  the  vesical  portion  has  been  taken  away;  s,  left  vesicula  seminalis  joining  the 
same  duct;  s,s,  the  right  ductus  deferens  and  right  vesicula  seminalis,  which  has  been 
unraveled;  />,  under  side  of  the  prostate  gland;  m,  part  of  the  urethra;  u,  u,  the  ureters 
(cut  short),  the  right  one  turned  aside.  (Haller.) 


THE    SEMINAL   FLUID 


771 


Head 


Body 


Tail 


enlargement  just  before  it  unites  with  its  fellow.  The  ductus  deferens  has 
muscular  walls  and  is  lined  with  ciliated  epithelial  cells. 

The  Vesiculae  Seminales. — The  seminal  vesicles  have  the  appearance 
of  outgrowths  from  the  base  of  the  deferent  ducts.  Each  deferent  duct 
just  before  it  enters  the  prostate  gland,  through  part  of  which  it  passes  to 
terminate  in  the  urethra,  gives  off  a  side  branch  which  bends  back  from  it  at 
an  acute  angle.  This  branch,  dilating,  variously  branching,  and  pursuing  in 
both  itself  and  its  branches  a  tortuous  course,  forms  the  vesicula  seminalis. 
Each  vesicle  is  a  single-branching,  convoluted,  and  sacculated  tube.  The 
microscopic  structure  resembles  closely  that  of  the  ductus  deferens. 

The  Penis. — The  penis  is  attached  to  the  symphysis  pubis  by  its  root. 
It  is  composed  of  three  long,  more  or  less  cylindrical  masses  enclosed  in 
remarkably  firm  fibrous  sheaths.  Two,  the 
corpora  cavernosa,  are  alike  and  are  firmly 
joined  together.  They  receive  below  and  be- 
tween them  the  third  part,  or  corpus  spongi- 
osum.  The  urethra  passes  through  the  corpus 
spongiosum.  The  enlarged  extremity,  or  glans 
penis,  is  continuous  with  the  corpus  spongiosum. 
Cowper's  glands  are  at  its  base,  and  their  ducts 
open  into  the  base  of  the  urethra. 

The  Prostate  Gland.— The  prostate  is 
situated  at  the  neck  of  the  urinary  bladder,  and 
encloses  the  base  of  the  urethra.  The  prostate 
is  made  up  of  small  compound  tubular  glands 
embedded  in  an  abundance  of  muscular  fibers 
and  connective  tissue.  The  glandular  sub- 
stance consists  of  numerous  small  saccules, 
opening  into  elongated  ducts,  which  unite  into 
a  smaller  number  of  excretory  ducts.  The 
acini  of  the  upper  part  of  the  prostate  are 
small  and  hemispherical,  in  the  middle  and 
lower  parts  the  tubes  are  longer  and  more 

convoluted.  The  ducts,  twelve  to  twenty  in  number,  open  into  the  urethra. 
They  are  lined  by  a  layer  of  columnar  cells,  beneath  which  is  a  layer  of 
small  polyhedral  cells. 

The  muscular  tissue  of  the  prostate  not  only  forms  the  chief  part  of  the 
stroma  of  the  gland,  but  also  forms  a  continuous  layer  inside  the  fibrous 
sheath,  as  well  as  a  layer  surrounding  the  urethra  continuous  with  the  sphinc- 
ter of  the  bladder. 

The  Seminal  Fluid. — The  sperm  cells  of  the  testes  are  joined  on  their 
way  to  the  exterior  by  the  fluids  secreted  by  the  mucous  lining  of  the  various 
tubules  and  glands.  Of  the  fluids  the  chief  ones  are  the  secretions  of  the 


End  piece — 


FIG.  491. — Human  Sperma- 
tozoa (after  Retzius).  A,  Side 
view,  B,  front  view. 


772  THE   REPRODUCTIVE    ORGANS 

seminal  vesicles,  of  the  prostate  gland,  and  of  Cowper's  glands.  The  sperm 
cells  and  the  secretions  together  constitute  the  seminal  fluid. 

After  the  period  of  puberty  the  seminal  fluid  is  secreted  constantly  but 
slowly,  except  under  sexual  excitement.  It  is  ordinarily  received  into  the 
seminal  vesicles,  whence  it  is  expelled  at  the  time  of  coitus.  In  celibates  the 
seminal  fluid  may  at  times  escape  in  small  quantity  into  the  urethra  to  be 
washed  away  by  the  urine,  or  periodic  reflex  emissions  may  occur.  The 
seminal  vesicles  contribute  a  secretion,  as  well  as  a  vesicle  to  receive  the 
sperm. 

The  secretion  of  the  seminal  vesicles  and  that  of  the  prostate  gland  are 
in  some  way  concerned  in  maintaining  the  activity  and  prolonging  the  life  of 
the  spermatozoa  probably  owing  to  the  alkalinity  of  the  secretions.  These 
cells  remain  alive  in  the  fluid  for  as  much  as  forty-eight  hours  after  removal 
from  the  body,  and  remain  alive  quite  indefinitely  in  the  vesicles  in  the  body. 
The  secretions  have  been  proven  necessary  to  the  life  and  function  of  the 
spermatozoa  by  the  results  of  operations  in  which  the  seminal  vesicles  and 
the  prostate  were  removed,  whereby  the  animals  became  sterile. 

THE  REPRODUCTIVE  ORGANS  OF  THE  FEMALE. 

The  female  genital  organs  consist  of  the  Ovarium,  the  Tuba  Uterina,  the 
Uterus,  and  the  Vagina. 


FIG.  492. — Diagrammatic  View  of  the  Uterus  and  Its  Appendages,  as  Seen  from 
Behind.  The  uterus  and  upper  part  of  the  vagina  have  been  laid  open  by  removing  the 
posterior  wall;  the  Fallopian  tube,  round  ligament,  and  ovarian  ligament  have  been  cut 
short,  and  the  broad  ligament  removed  on  the  left  side,  u,  The  upper  part  of  the  uterus; 
c,  the  cervix  opposite  the  os  internum;  the  triangular  shape  of  the  uterine  cavity  is  shown, 
and  the  dilatation  of  the  cervical  cavity  with  the  rugae  termed  arbor  vitae;  v,  upper  part  of 
the  vagina;  od,  Fallopian  tube  or  oviduct;  the  narrow  communication  of  its  cavity  with  that 
of  the  cornu  of  the  uterus  on  each  side  is  seen;  I,  round  ligament;  lo,  ligament  of  the  ovary; 
o,  ovary;  i,  wide  outer  part  of  the  right  Fallopian  tube;  fiy  its  fimbriated  extremity;  PO, 

rrovarium;  h,  one  of  the  hydatids  frequently  found  connected  with  the  broad  ligament. 
(Allen  Thomson.) 

The  Ovaries. — The  ovaries  are  paired  bodies,  situated  in  the  cavity  of 
the  pelvis,  and  adherent  to  the  posterior  surface  of  the  broad  ligament.  The 


THE    OVARIES 


773 


attached  border  of  the  ovary  is  called  the  hilum,  and  it  is  at  this  point  that  the 
blood  vessels  and  nerves  enter  it.  Each  ovary  is  about  4  cm.  long,  2  cm. 
wide,  and  1.25  cm.  thick.  It  is  supported  by  the  suspensory  ligament. 

The  internal  structure  of  the  ovary  in  all  mammals  consists  of  a  peculiar 
soft  fibrous  connective  tissue,  stroma,  abundantly  supplied  with  blood  vessels. 
The  surface  of  the  ovary  is  covered  with  cubical  epithelium.  Embedded  in 
the  stroma  in  various  stages  of  development  are  numerous  minute  follicles 
or  vesicles,  the  vesicular  ovarian  follicles,  containing  the  ova,  figure  494. 
They  are  small  and  numerous  near  the  surface  of  the  ovary,  either  arranged 
as  a  continuous  layer,  as  in  the  cat  or  rabbit,  or  in  groups,  as  in  the  human 
ovary.  Nearer  the  center  are  large  and  fully  developed  follicles. 

Each  follicle  has  an  external  membranous  envelope,  or  tunica  externa, 
which  is  lined  with  a  layer  of  nucleated  cells,  forming  a  kind  of  epithelium 


FIG.  493. — Diagrammatic  Section  of  the  Ovary,  Showing  its  Cortical  or  Ovigenous 
Layer,  Formed  of  Ovisacs  in  Various  Stages  of  Evolution.  (Duval.)  A,  A,  A,  Primordial 
ovisacs;  B,  B,  B,  ovisacs  further  developed;  C,  ovisac  approaching  maturity;  D,  ripe  ovisac 
with  its  proligerous  disc  (DP)  containing  the  ovum;  MG,  membrana  granulosa;  H,  hilum 
of  ovary. 

or  internal  coat  and  named  the  tunica  interna.  The  cavity  of  the  follicle  con- 
tains the  ovule,  or  immature  egg  cell,  enclosed  in  a  very  delicate  membrane. 
The  large  spherical  nucleus  contains  one  or  more  nucleoli.  The  nucleus  is 
known  as  the  germinal  vesicle,  and  the  nucleolus  as  the  germinal  spot. 

The  human  ovum  measures  about  0.2  mm.  in  diameter.  Its  external 
investment,  or  the  zona  pellucida,  or  vitelline  membrane,  is  a  transparent 
membrane,  about  io/*  in  thickness,  which  under  the  microscope  appears 
as  a  bright  ring,  figure  495.  The  ovum  itself  has  the  characteristic  structure 
of  the  typical  cell,  with  the  exception  that  its  cytoplasm  is  filled  with  nu- 
merous yolk  granules.  The  larger  granules  or  globules,  which  have  the 
aspect  of  fat-globules,  are  in  greatest  number  at  the  periphery  of  the  yolk. 

The  nucleus,  or  germinal  vesicle,  is  about  o .  05  mm.  in  diameter.  The 
vesicle  is  of  greatest  relative  size  in  the  smallest  ova. 


774 


THE    REPRODUCTIVE    ORGANS 


These  ova  are  descended  from  primitive  germ  cells  which  become  differ- 
entiated very  early  in  the  embryo.  In  some  vertebrates,  such  as  the  chick, 
there  is  evidence  that  the  primitive  germ  cells  become  distinct  from  all  other 
cells  in  the  body  even  before  the  formation  of  the  embryonic  mesoderm  has 
been  completed.  Later  they  become  located  in  a  membrane  of  short  colum- 
nar cells,  the  so-called  germinal  epithelium,  covering  the  surface  of  the  embry- 
onic ovary. 

The  Graafian  follicles  of  the  human  ovary  are  formed  in  the  following 
manner:  The  cells  of  the  germinal  epithelium  undergo  proliferation  so  as 
to  form  several  strata,  and  grow  into  the  ovarian  stroma  as  longer  or  shorter 


Downgrowths  of  epithelium         Ovum  with  its  investing  cells 
Germinal  epithelium 


Stratum  granulosum 


Epithelial  cells        Ovarian  strorna  Graafian  follicle 


Ovum 


Liquor  folliculi 
Discus  proligerus 


FIG.  494. — A,  Diagrammatic  Representation  of  the  Manner  in  which  the  Vesicular 
Ovarian  Follicles  Arise  During  the  Development  of  the  Ovary.  B,  Diagram  Illustrating 
the  Structure  of  a  Ripe  Vesicular  Ovarian  Follicle.  (Cunningham.) 


columns  or  tubes.  By  degrees  these  tubes  become  cut  off  from  the  surface 
epithelium,  and  form  cell  nests,  small  if  near  the  surface,  larger  if  in  the  depth 
of  the  stroma.  The  nests  increase  in  size  from  multiplication  of  their  cells. 
Inside  these  nests,  certain  cells  which  are  descended  from  the  primitive  germ 
cells  enlarge  and  form  ova.  The  small  cells  of  a  nest  surround  the  ova,  and 
form  their  membrana  granulosa,  and  the  stroma  growing  up  separates  the 
surrounded  ova  into  so  many  Graafian  follicles. 

The  smallest  follicles  are  formed  at  the  surface,  and  make  up  the  cortical 
layer.  It  is  said  by  some  that  the  superficial  follicles  as  they  begin  to  ripen 
become  more  deeply  placed  in  the  ovarian  stroma;  and,  again,  that  as  they 
increase  in  size  they  make  their  way  back  toward  the  surface.  The  develop- 
ment of  all  eggs  that  are  destined  to  mature  is  carried  as  far  as  the  early  fol- 
licular  stage  in  the  ovary  of  the  child,  previous  to  the  birth  of  the  child,  or 
within  a  relatively  short  period  thereafter.  Conditions  indicative  of  the  for- 


THE    UTERINE   TUBES   OR   OVIDUCTS 


775 


mation  of  new  follicles  have  never  been  observed  in  a  person  more  than  two 
years  of  age. 

When  the  vesicular  ovarian  follicles  mature,  they  form  little  prominences 
on  the  exterior  of  the  ovary  covered  only  by  a  thin  layer  of  condensed  fibrous 
tissue  and  epithelium.  From  the  earliest  infancy,  and  through  the  whole 
fruitful  period  of  life,  there  appears  to  be  a  constant  development  and  matur- 
ing of  ovarian  vesicles,  with  their  contained  ova.  Until  the  period  of  puberty, 
however,  the  process  is  comparatively  inactive.  But,  coincident  with  the 


FIG.  495. — Diagrammatic.  Representation  of  a  Human  Ovum  and  Its  Coverings. 
(Cunningham.) 

The  corona  radiata,  which  completely  surrounds  the  ovum,  is  represented  only  in  the 
lower  part  of  the  figure. 

1,  Corona  radiata;  5,  vitellus  or  yolk; 

2,  granular  layer;  6,  germinal  vesicle  (nucleus); 

3,  vitelline  membrane;  7,  germinal  spot  (nucleolus); 

4,  zona  pellucida  (oolemma);  8,  nuclear  membrane. 


i 


other  changes  which  occur  in  the  body  at  the  time  of  puberty,  the  ovaries 
enlarge  and  become  very  vascular,  the  development  of  ovarian  vesicles  is  more 
abundant,  the  size  and  degree  of  development  attained  by  them  are  greater, 
and  the  ova  are  capable  of  being  fertilized.  The  follicles  receive  their 
nourishment  from  the  surrounding  capillaries  in  the  stroma.  They  are  never 
penetrated  by  blood  vessels,  and  are  never  entered  by  nerve  fibers,  so  far  as  is 
known. 

The  Uterine  Tubes  or  Oviducts. — The  uterine  tubes  are  about  10 
cm.  in  length  and  extend  between  the  ovaries  and  the  upper  angles  of  the 
uterus.  At  the  point  of  attachment  to  the  uterus  each  tube  is  very  narrow; 
but  in  its  course  to  the  ovary  it  increases  to  about  3  mm.  in  thickness.  At 


THE    REPRODUCTIVE    ORGANS 

its  distal  extremity,  which  is  free  and  floating,  it  bears  a  number  of  elongated 
lobes,  or  fimbrice,  one  of  which  is  longer  than  the  rest  and  is  attached  to  the 
ovary.  The  canal  of  the  tube  is  narrow,  especially  at  its  point  of  entrance 
into  the  uterus.  Its  other  extremity  is  wider  and  opens  into  the  cavity  of 
the  abdomen  at  the  fimbriae.  The  uterine  tube  is  invested  with  peritoneum, 
and  its  canal  is  lined  with  ciliated  epithelium. 

The  Uterus. — The  uterus,  u,  c,  figure  492,  is  a  somewhat  pear  shaped 
organ,  and  is  about  7.5  cm.  in  length,  5  cm.  in  breadth  at  its  upper  part 
or  fundus,  but  at  the  neck  or  cervix  only  about  1.25  cm.  The  part  be- 
tween the  fundus  and  neck  is  termed  the  body  of  the  uterus;  it  is  about 
2.5  cm.  in  thickness. 

The  uterus  is  constructed  of  three  principal  layers,  or  coats:  serous, 
fibrous  and  muscular,  and  mucous.  The  serous  coat,  which  has  the  same 
general  structure  as  the  peritoneum,  covers  the  organ  except  the  front  surface 
of  the  neck.  The  middle  coat  is  a  thick  mass  of  unstriped  muscle.  The 
muscle  fibers  become  enormously  developed  during  pregnancy.  The  arteries 
and  veins  are  found  in  large  numbers  in  the  outer  part  so  as  to  form  almost  a 
special  vascular  coat.  The  mucous  membrane  of  the  uterus  is  composed  of 
columnar  ciliated  epithelium,  which  extends  also  to  the  interior  of  the 
tubular  glands,  of  which  the  mucous  membrane  is  largely  made  up.  In  the 
cervix  of  the  uterus  the  mucous  membrane  is  arranged  in  permanent  longi- 
tudinal folds,  pliccB  palmate.  The  glands  of  this  part  branch  repeatedly, 
and  extend  deeply  into  the  substance  of  the  cervix.  The  body  has  numerous 
simpler  tubular  glands.  The  glands  are  also  lined  with  ciliated  epithelium. 
They  secrete  a  thick  glairy  mucus,  resembling  white  of  egg. 

The  Vagina. — The  vagina  is  a  membranous  canal  8  to  10  cm.  long, 
extending  obliquely  downward  and  forward  from  the  neck  of  the  uterus, 
which  it  embraces,  to  the  external  organ  of  generation.  It  is  lined  with 
mucous  membrane,  covered  with  stratified  squamous  epithelium,  which  in 
the  ordinary  contracted  state  of  the  canal  is  thrown  into  transverse  folds. 
External  to  the  mucous  membrane,  the  walls  of  the  vagina  are  constructed 
of  unstriped  muscle  and  fibrous  tissue,  within  which  in  the  submucosa, 
especially  around  the  lower  part  of  the  tube,  is  a  layer  of  erectile  tissue.  The 
lower  extremity  of  the  vagina  is  embraced  by  an  orbicular  muscle,  the 
sphincter  vagina.  The  external  organs  of  generation  are  the  clitoris,  the 
Idbia  interna  or  nymphce;  and,  the  labia  externa  or  pudenda,  formed  of  the 
external  integument,  and  lined  internally  by  mucous  membrane.  Numerous 
mucous  follicles  are  scattered  beneath  the  mucous  membrane  of  the  external 
organs  of  generation;  and  two  larger  lobulated  glands,  the  glands  of  Ear- 
th olin,  analogous  to  Cowper's  glands  in  the  male,  are  located  at  the  sides 
of  the  lower  part  of  the  vagina.  The  ducts  of  these  glands  are  about  12  mm. 
long  and  open  immediately  external  to  the  hymen  at  the  mid- point  of  the 
lateral  wall  of  the  vaginal  orifice. 


OVULATION    AND    MENSTRUATION 


777 


Ovulation  and  Menstruation. — In  the  process  of  development  in  the 
ovary,  the  individual  vesicular  ovarian  follicle  increases  in  size  and  gradually 
approaches  the  surface  of  the  ovary.  When  fully  ripe  or  mature,  it  forms  a 
little  projection  on  the  exterior.  Coincident  with  the  increase  in  size,  which 
is  caused  by  the  augmentation  of  its  liquid  contents,  the  external  envelope 
of  the  distended  vesicle  becomes  very  thin  and  eventually  bursts.  The  ovum 
and  fluid  contents  of  the  vesicle  escape  on  the  exterior  of  the  ovary,  whence 
they  pass  into  the  uterine  tube. 


FIG.  496. 


FIG.  497. 


FIG.  498. 


FIG.    496. — Diagram   of  Uterus   just   Before   Menstruation.     The   shaded   portion 

represents  the  thickened  mucous  membrane. 
FIG.  497. — Diagram  of  Uterus  when  Menstruation  has  just  Ceased,  Showing  the 

Cavity  of  the  Uterus  Deprived  of  Mucous  Membrane. 

FIG.  498.— Diagram  of  Uterus  a  Week  After  the  Menstrual  Flux  has  Ceased.     The 
shaded  portion  represents  renewed  mucous  membrane.     (J.  Williams.) 

In  man  and  mammals  ovulation  apparently  occurs  only  at  certain  periods. 
These  periods  are  closely  connected  with  the  changes  in  the  woman  that  con- 
stitute the  phenomenon  of  menstruation,  or,  in  the  lower  mammals,  of  oestrus, 
or  heat. 

That  ovulation  and  discharge  occur  periodically,  and  only  during  the 
phenomenon  of  heat,  in  the  lower  mammalia,  is  indicated  by  the  facts 
that,  in  all  instances  in  which  ovarian  vesicles  have  been  found  presenting 
the  appearance  of  recent  rupture,  the  animals  were  at  the  time  or  had  recently 
been  in  heat.  There  are  few  authentic  and  detailed  accounts  of  ovarian 
vesicles  being  found  ruptured  or  of  ova  being  fertilized  in  the  intervals 


778  THE    REPRODUCTIVE    ORGANS 

between  periods  of  heat.  Although  conception  is  not  confined  to  the  periods 
of  menstruation  in  the  human  species,  yet  it  is  more  likely  to  occur  about  a 
menstrual  epoch  than  at  other  times. 

The  exact  relation  between  the  discharge  of  ova  and  menstruation  is  not 
very  clear.  In  animals,  the  physiological  analogy  of  menstruation  appears  to 
be  found  in  the  proaestral  processes,  which  are  the  preliminary  stages  of  heat, 
and  occur  immediately  before  ovulation.  It  was  formerly  believed  that  men- 
struation was  the  result  of  a  congestion  of  the  uterus  arising  in  association 
with  the  enlargement  and  rupture  of  a  vesicular  ovarian  follicle;  but  though  a 
vesicular  ovarian  follicle  is,  as  a  rule,  ruptured  at  each  menstrual  epoch,  yet 
instances  are  recorded  in  which  menstruation  has  occurred  where  no  ovarian 
follicle  can  have  been  ruptured,  and  cases  where  ova  have  been  discharged  in 
amenorrheic  women.  It  must  therefore  be  admitted  that  menstruation  is 
not  strictly  dependent  on  the  maturation  and  discharge  of  ova. 

Observations  made  after  death,  and  facts  obtained  by  clinical  investiga- 
tion, support  the  view  that  rupture  of  a  vesicular  ovarian  follicle  does  not 
happen  on  the  same  day  of  the  monthly  period  in  all  women.  In  the  minor- 
ity of  cases  it  may  occur  toward  the  close  or  soon  after  the  cessation  of  a 
flow.  On  the  other  hand,  in  almost  all  subjects  examined  after  death,  of 
which  there  is  record,  rupture  of  the  follicle  appears  to  have  taken  place 
before  the  commencement  of  the  menstrual  flow. 

However,  the  presence  of  the  ovaries  seems  necessary  for  the  performance 
of  the  menstrual  function;  for  women  do  not  menstruate  when  both  ovaries 
have  been  removed  by  operation.  See  page  498  for  a  discussion  of  the 
functional  effects  of  removal  of  the  ovary. 

Source  and  Character  of  Menstrual  Changes. — The  menstrual  periods 
usually  occur  at  intervals  of  a  lunar  month,  the  duration  of  each  being  from 
three  to  six  days.  In  some  women  the  intervals  are  so  short  as  three  weeks 
or  even  less;  while  in  others  they  are  longer  than  a  month.  The  periodical 
return  is  usually  attended  by  pains  in  the  loins,  a  sense  of  fatigue  in  the  lower 
limbs,  and  other  symptoms,  which  vary  extremely  in  different  individuals. 

The  menstrual  discharge  is  a  thin  sanguineous  fluid,  and  consists  of  blood, 
epithelium,  and  mucus  from  the  uterus  and  vagina.  The  menstrual  flow 
is  preceded  by  a  general  engorgement  of  all  the  pelvic  organs  with  blood. 
The  cervix  and  vagina  become  darker  in  color  and  softer  in  texture,  and 
the  quantity  of  mucus  secreted  by  the  glands  of  the  cervix  and  body  is  in- 
creased. The  uterine  mucous  membrane  is  swollen  and  the  glands  are 
enlarged.  The  discharge  of  blood,  the  source  of  which  is  the  mucous 
membrane  of  the  body  of  the  uterus,  is  probably  associated  with  uterine 
contractions.  There  is  great  difference  of  opinion  as  to  whether  or  not 
any  of  the  uterine  mucous  membrane  is  normally  shed  during  the  process 
of  menstruation.  John  Williams  believes  that  the  whole  of  the  mucous 
membranes  of  the  body  of  the  uterus  is  thrown  off  at  each  monthly  period, 


MENSTRUAL   LIFE  779 

forming  a  true  decidua  menstrualis,  figure  496,  while  Moricke  and  others 
believe  that  the  mucous  membrane  remains  intact.  Leopold  believes 
that  red  blood  corpuscles  escape  from  the  congested  capillaries  and  under- 
mine the  superficial  epithelium,  and  that  in  this  way  the  superficial  layer  of 
the  mucous  membrane  is  eroded  and  subsequently  regenerated.  There 
is  a  period  of  regeneration  followed  by  a  period  of  rest  before  the  next 
repetition.  Minot  distributes  the  variations  in  time  as  follows: 

Tumefaction 5  days 

Menstrual  discharge 4  days 

Restoration  of  mucosa 7  days 

Period  of  rest 12  days 

The  menstrual  period  is  often  accompanied  by  profound  disturbances  in 
other  parts  of  the  body,  especially  of  the  vascular  and  of  the  nervous  systems, 
and  of  the  nutritive  processes. 

CorpusLuteum. — Immediately  before,  as  well  as  subsequent  to,  the  rupture 
of  an  ovarian  follicle  and  the  escape  of  its  ovum,  changes  ensue  in  the  interior 
of  the  follicle,  which  result  in  the  production  of  a  yellowish  mass,  termed  a 
corpus  luteum. 

When  fully  formed,  the  corpus  luteum  of  mammals  is  a  roundish  solid 
body,  of  a  yellowish  or  orange  color,  and  composed  of  a  number  of  lobules, 
which  surround,  sometimes  a  small  cavity,  but  more  frequently  a  small 
stelliform  mass  of  substance,  from  which  delicate  processes  pass  as  septa 
between  the  several  lobules.  The  processes  gradually  change  till  they 
nearly  fill  the  cavity  of  the  follicle,  and  even  protrude  from  the  orifice  in 
the  external  covering  of  the  ovary.  Subsequently  this  orifice  closes,  but  the 
fleshy  growth  within  still  increases  during  the  earlier  period  of  pregnancy, 
the  color  of  the  substance  gradually  changing  to  yellow,  and  its  consistency 
becoming  firmer.  After  the  orifice  of  the  follicle  has  closed,  the  growth 
of  the  yellow  substance  continues  during  the  first  half  of  pregnancy,  till 
the  cavity  is  reduced  to  a  comparatively  small  size  or  is  obliterated;  in  the 
latter  case,  merely  a  white  stelliform  cicatrix  remains  in  the  center  of  the 
corpus  luteum. 

The  first  changes  of  the  internal  coat  of  the  ovarian  follicle  in  the  proc- 
ess of  formation  of  a  corpus  luteum  seem  to  occur  in  every  case  in  which  an 
ovum  escapes.  If  the  ovum  is  impregnated,  the  growth  of  the  yellow  sub- 
stance continues  during  nearly  the  whole  period  of  gestation  and  forms  the 
large  corpus  luteum  commonly  described  as  a  characteristic  mark  of 
impregnation. 

The  significance  of  the  corpus  luteum  is  found  in  the  belief  that  it  is  the 
portion  of  the  ovary  especially  concerned  in  the  production  of  an  internal 
secretion  that  affects  the  uterus,  especially  stimulating  it  at  and  before  the 
menstrual  period. 


780  THE   REPRODUCTIVE    ORGANS 

Menstrual  Life. — The  occurrence  of  a  menstrual  discharge  is  one  of 
the  most  prominent  indications  of  the  commencement  of  puberty  in  the  fe- 
male sex;  though  its  absence  even  for  several  years  is  not  necessarily  at- 
tended with  arrest  of  the  other  characters  of  this  period  of  life  or  incapability 
of  impregnation.  The  average  time  of  its  first  appearance  in  females  of 
this  country  and  others  of  about  the  same  latitude  is  from  fourteen  to  fifteen: 
but  it  is  much  influenced  by  the  kind  of  life  to  which  girls  are  subjected, 
being  accelerated  by  habits  of  luxury  and  indolence,  and  retarded  by  con- 
trary conditions.  Its  appearance  may  be  slightly  earlier  in  persons  dwelling 
in  warm  climates  than  in  those  inhabiting  colder  latitudes.  The  menstrual 
functions  continue  through  the  whole  fruitful  period  of  a  woman's  life,  and 
usually  cease  between  the  forty-fifth  and  fiftieth  years,  which  time  is  known 
as  the  climacteric.  Menstruation  does  not  usually  occur  in  pregnant  women. 


CHAPTER  XVII. 
DEVELOPMENT. 

Maturation  of  Male  Germ  Cells,  the  Spermatozoon. — The  general 
effect  of  maturation  of  germinal  cells  is  to  reduce  the  number  of  chromo- 
somes, in  order  that  there  may  not  be  more  than  the  normal  supply  in  the 
new  cell  that  will  be  formed  by  the  union  of  the  ovum  and  the  sperm 
in  the  fertilization  of  the  ovum.  We  shall  describe  the  process  of 
maturation  in  terms  that  will  apply  to  the  production  of  spermatozoa 
in  nearly  all  animals. 

In  each  species  of  animals  the  number  of  chromosomes  in  every  cell 
nucleus  is  constant.  Although  they  are  of  various  sizes  and  shapes,  each 
nucleus  in  the  body  appears  to  have  the  same  assortment  as  every  other.  This 
is  explained  by  the  fact  that  whenever  a  cell  divides,  each  chromosome  divides 
individually,  and  the  two  resulting  daughter  chromosomes  from  each  original 
chromosome  are  distributed  to  the  two  new  cells.  Thus  the  chromosomes 
can  be  traced  back  individually  through  innumerable  cell  divisions  to  the 
assortment  that  existed  in  the  egg  cell  at  the  very  beginning  of  its  development. 
The  source  of  this  supply  of  chromosomes  found  in  the  developing  egg  is 
double,  one  set  having  been  derived  from  the  original  nucleus  of  the  egg,  and 
the  other  provided  by  the  sperm.  Now  the  developed  organism,  which  has 
obtained  in  this  way  a  double  set  of  chromosomes  in  every  cell,  must  like  its 
ancestors  be  able  to  produce  germ  cells  that  each  contain  a  single  set. 

This  maturation  reduction  of  chromosomes,  from  a  double  to  a  single  set, 
is  always  carried  out  by  the  joint  effect  of  a  process  of  synapsis  and  of  two 
peculiar  cell  divisions  known  as  the  maturation  divisions.  In  the  nucleus  of 
the  primary  spermatocyte  the  chromatin  materials  become  crowded  together, 
and  pass  through  the  stage  known  as  synapsis,  during  which  the  chromosomes 
become  closely  united  with  each  other  in  pairs.  The  double  set  of  simple,  i.e., 
uncompounded  chromosomes  occurring  in  the  spermatogonia  is  thus 
transformed  in  the  primary  spermatocyte  into  an  apparently  single  set  of 
double  chromosomes.  Each  of  these  double  structures  is  believed  to  repre- 
sent the  union  of  corresponding  chromosomes  out  of  the  two  sets  previously 
existing.  They  grow  immediately  into  a  four-parted  condition,  which  has 
given  them  the  name  tetrads.  In  the  cell  division  of  the  primary  spermato- 
cytes,  the  first  maturation  division,  the  tetrads  are  divided  equally,  and  the 
result  is  two  secondary  spermatocytes,  each  containing  a  single  set,  called 
diads.  The  second  maturation  division  follows  promptly,  producing  two 

781 


782 


DEVELOPMENT 


spermatids  from  each  secondary  spermatocyte.     Each  spermatid  receives 
half  the  chromosomes  of  each  diad,  and  as  a  result  a  spermatid  possesses  but 


Spermatozoa 


FlG.  499. 

a  single  set  of  simple  chromosomes.     It  is  then  said  to  have  the  reduced  or 
gametic  number,  characteristic  of  sexual  germ  ceils,  as  contrasted  with  the 


FIG.  500. 

FIGS.  499  and  500. — Diagrams  Showing  the  Scheme  of  Development  and  Chromatin 
Reduction  in  the  Growth  of  the  Spermatozoa  and  Ova.     (From  Cunningham.) 

unreduced  or  somatic  number,  characteristic  of  the  cells  of  the  body. 

The  spermatid  is  metamorphosed  directly  into  a  spermatozoon.     The 


MATURATION   OF    THE   OVUM  — THE   FEMALE    GERM   CELL 


783 


process  consists  in  the  development  of  a  tail  or  cilium,  the  transformation  of 
the  nucleus  into  the  form  of  a  spermatozoon  head,  and  the  sloughing  off 
of  nearly  all  the  remaining  cell  protoplasm.  The  centrosome,  a  minute  body 
closely  concerned  with  the  power  of  cell  division,  is  apparently  retained,  but 
the  power  of  cell  division  is  impossible  in  the  presence  of  so  little  cytoplasm, 
figure  499. 


FIG.  501.— The  Maturation  of  the  Ovum;  Extrusion  of  the  "Polar  Bodies."  (Dia- 
grammatic.) A,  An  ovum  at  the  commencement  of  the  process;  B,  after  the  formation  of 
the  spindle.  The  chromosomes  are  gathered  at  the  equator  of  the  spindle.  C,  One  apex 
of  the  spindle  has  projected  into  a  bud  on  the  surface,  and  half  of  the  divided  dyads  have 
passed  to  each  pole;  D,  the  separation  of  the  first  polar  body;  E,  the  commencement  of  the 
second  polar  body;  F,  the  completion  of  the  second  polar  body.  (Cunningham.) 

Maturation  of  the  Ovum — the  Female  Germ  Cell. — The  ovum  when 
liberated  from  the  ovary  is  a  single  cell  enclosed  within  the  zona  pellucida, 
and  containing  the  germinal  vesicle  or  nucleus  and  germinal  spot  or  nucleolus. 
It  still  possesses  the  somatic  or  double  number  of  chromosomes,  and  must 
undergo  maturation  divisions  in  order  to  reduce  their  numbers.  Previous 
to  the  maturation  divisions  it  should  more  correctly  be  called  a  primary 
oocyte. 

Synapsis  now  occurs  in  the  nucleus.  It  is  not  fundamentally  different 
from  the  synapsis  in  the  spermatocyte.  Meanwhile  the  nucleus  migrates 
to  the  surface  of  the  oocyte.  After  synapsis,  the  chromosomes  are  in  the  form 
of  a  single  set  of  tetrads  as  in  the  male.  They  are  liberated  by  the  disappear- 


784  DEVELOPMENT 

ance  of  the  nuclear  membrane,  and  become  arranged  upon  a  nuclear  spindle, 
as  if  for  an  ordinary  cell  division.  The  two  maturation  divisions  occur  in 
rapid  succession.  In  the  first  of  these  the  oocyte  is  divided  extremely  un- 
equally, into  a  secondary  occyte  of  nearly  the  same  size  as  the  primary,  and  the 
first  polar  body,  morphologically  equivalent  to  a  minute,  degenerant  secondary 
oocyte.  Each  secondary  oocyte  receives  a  single  set  of  diads,  which  are  the 
division  products  of  the  set  of  tetrads.  The  second  maturation  division  is 
similarly  unequal,  and  its  products  are  the  ripe  ovum  and  the  second  polar 
body.  Each  of  these  receives  a  single  set  of  simple  chromosomes.  Those 
that  are  in  the  ovum  become  the  female  pronucleus,  possessing  the  reduced 
chromosomal  number,  figure  500.  The  centrosome  disappears,  and  the  egg 
is  left  incapable  of  division.  It  remains  unchanged  unless  fertilized  by  the 
sperm. 

Material  Basis  of  Heredity. — In  considering  the  relations  of  parent  and 
child  to  each  other  through  heredity,  it  is  necessary  to  distinguish  sharply 
between  that  which  comes  to  the  child  as  an  inherent  part  of  its  nature,  and 
all  else  which  may  be  added  through  conditions  of  nourishment,  physical 
surroundings,  infection  with  disease-producing  parasites,  and  the  like. 
The  former  is  the  only  true  heredity,  in  a  scientific  sense,  although  it  is 
obvious  that  until  birth,  or  even  for  some  time  after  birth,  the  well  being  and 
development  of  the  germ  cells,  fetus,  and  child  are  directly  dependent  upon 
the  parental  organism. 

It  has  been  shown  that  every  cell  in  a  child  is  descended  from  one  cell,  the 
fertilized  ovum.  This  fertilized  egg  is  the  product  of  the  union  of  male  and 
female  germ  cells,  cells  which  are  not  derived  from  the  active  body  tissues  of 
the  parents,  but  can  be  traced  back  to  cells  that  were  set  aside  for  the  purpose 
of  reproduction  at  a  very  early  stage  of  the  parent's  embryonic  life.  There- 
after the  influences  which  the  vicissitudes  of  the  parent's  life  may  have  exerted 
upon  the  germ  cells  he  produces  are  limited  to  what  can  be  carried  by  diffu- 
sion into  the  germ  cells  of  substances  from  the  lymph. 

Relative  Influence  of  Mother  and  Father  in  Heredity. — We  may  use  a 
more  concrete  form  of  this  question  as  follows:  If  two  races  are  crossed,  does 
it  make  any  difference  which  form  was  the  mother's  stock,  and  which  the 
father's  ?  This  comparison  has  been  made  very  extensively  in  domesticated 
animals  and  garden  plants,  and  to  a  less  extent  in  man.  Good  examples  are 
the  inheritance  of  color  peculiarities  in  mammals,  and  peculiarities  of  the 
skeleton.  Excepting  in  certain  special  cases,  like  color  blindness  in  man, 
which  will  be  treated  later,  it  is  found  that  these  "reciprocal  crosses"  pro- 
duce identical  results  in  heredity;  hence,  that  the  average  influence  of  father 
and  mother  upon  inheritance  is  in  general  equal.  We  have  already  seen 
that  the  two  sexes  make  very  unequal  contributions  to  the  substance  of 
the  fertilized  egg.  The  following  table  summarizes  these  differences 
and  similarities. 


MENDELIAN    INHERITANCE    AND    CHROMOSOMES  785 

Ripe  ovum  Much  cytoplasm         Single  set  of  chromosomes       No  centrosome 

Spermatozoon          Trace  of  cytoplasm    Single  set  of  chromosomes       One  centrosome 

Fertilized  ovum        Much  cytoplasm        Double  set  of  chromosomes     One  centrosome 

We  see  that  virtually  all  the  cytoplasm  of  the  beginning  embryo  comes  from 
the  mother,  and  represents  her  variety.  The  minute  centrosome  comes  from 
the  spermatozoon.  But  the  nucleus  represents  the  two  stocks  half  and  half, 
and  is  the  only  part  of  the  germ  cell  that  fulfills  the  conditions  for  equal  inheri- 
tance from  the  two  parental  stocks.  The  conclusion  has  been  drawn,  that 
the  transmission  of  such  differences  as  we  find  between  different  breeds  of  the 
same  species  is  due  to  peculiarities  in  the  nuclei. 

Mendelian  Inheritance  and  Chromosomes. — Two  types  of  inheritance 
have  long  been  recognized,  the  blending,  and  the  alternative  or  Mendelian 
forms.  In  the  former,  the  offsprings  are  intermediate  between  the  stocks  of 
their  parents.  In  the  latter,  a  certain  characteristic  from  one  stock  is  mani- 
fested by  the  child  to  the  exclusion  of  the  alternative  trait  shown  in  the  other 
stock.  Many  supposed  cases  of  blending  are  now  believed  to  be  complex 
instances  of  alternative  inheritance.  Albinism,  or  the  lack  of  pigment  in  hair, 
skin,  and  eyes,  is  a  good  example  of  an  alternatively  heritable  trait. 

When  two  pure  bred  animals  that  carry  two  opposite  and  alternative 
traits  are  crossed  with  each  other,  one  type  "dominates,"  or  shows  in  full 
force  in  all  the  young.  Thus  in  most  mammals  color  dominates  over  albin- 
ism, and  all  the  young  from  the  cross  look  like  thoroughbred  colored  animals. 
We  conclude  that  if  either  egg  or  sperm  carries  color,  the  offspring  is  always 
colored  in  appearance.  But  if  these  animals  are  now  crossed  with  the  white 
stock,  half  of  the  offspring  will  be  white  and  the  others  colored.  This  indi- 
cates that  the  dark  colored  hybrids  are  producing  equal  numbers  of  two  kinds 
of  germ  cells,  one  of  which  possesses,  and  the  other  lacks  the  color-producing 
factor.  These  last  colored  young  are  really  hybrids  producing  two  kinds  of 
germ  cells,  like  their  hybrid  parent.  But  their  white  brothers  have  received 
white  from  both  their  parents,  and  are  essentially  a  pure  stock  once  more. 

Thus  in  simple  alternative  (Mendelian)  inheritance,  the  hybrid  looks  like 
a  pure-bred  representative  of  the  "dominant"  form,  the  other,  or  "recessive" 
character  not  coming  to  view.  And  its  germ  cells  are  not  hybrid,  but  half 
of  them  are  pure  for  one  form,  and  the  other  half  pure  for  the  other  form. 
The  character  of  the  offspring  depends  upon  the  particular  germ  cells  that 
produced  it.  If  they  are  both  alike,  the  young  is  "pure,"  no  matter  how 
hybrid  its  parents. 

These  facts  lead  to  certain  statistical  relations  known  as  the  Mendelian 
ratios,  in  the  proportionate  numbers  of  pure  dominant,  mixed,  and  pure 
recessive  young  that  will  be  found  in  each  kind  of  cross.  When  two  pairs  of 
Mendelian  characters  are  crossed  at  once,  as  for  example,  two  colors  of  hair 
and  two  contrasting  lengths  of  hair,  each  pair  of  characters  usually  follows 
these  Mendelian  rules  quite  independent  of  the  other  pair.  From  one  such 


786  DEVELOPMENT 

hybrid  animal,  germ  cells  will  result  that  carry  white  short  hair,  white  long, 
dark  short,  and  dark  long. 

The  theory  of  inheritance  through  the  agency  of  chromosomes  is  based 
upon  this  type  of  fact,  and  the  close  similarity  to  the  manner  of  distribution 
of  chromosomes  in  the  maturation  of  the  germ.  By  the  theory  the  color 
hybrid  is  supposed  to  have  a  color-producing  chromosome  from  one  parent, 
but  from  the  other  parent  a  corresponding  chromosome  which  lacks  this 
power.  In  the  maturation  of  the  germ  cell  one  of  these  two  corresponding 
chromatin  elements  is  left  alone  in  each  mature  cell,  otherwise  the 
chromosomes  would  not  have  been  reduced  from  a  double  to  a  single  set. 
And  for  any  one  germ  cell  the  chances  are  equal  whether  it  receives  the  color- 
bearing  or  the  color-lacking  chromosome.  In  either  case  the  cell  is  "pure" 
for  whichever  trait  it  carries. 

Germ  Cells  in  Relation  to  the  Determination  of  Sex. — A  peculiar 
chromosome  or  pair  of  chromosomes  has  been  found  in  man,  and  some  other 
vertebrates,  as  well  as  in  insects,  nematode  worms  and  other  types  of  lower 
animals,  which  is  closely  connected  with  the  determination  of  sex.  It  is 
called  the  heterochromosome,  or  X  chromosome.  In  its  simplest  form  this 
chromosome  is  single  in  the  male,  but  paired  in  the  female.  In  synapsis  in 
the  male  cells  it  remains  separate,  because  of  the  lack  of  a  mate.  As  a  result, 
before  the  first  maturation  division  it  is  a  triad  instead  of  a  tetrad.  It  can 
divide  only  once  in  the  two  maturation  divisions,  and  at  the  other  division  it 
passes  bodily  over  to  one  of  the  daughter  cells,  leaving  the  other  cell  unsup- 
plied.  Thus  there  are  produced  equal  numbers  of  two  types  of  spermatozoa, 
those  with  and  those  without  an  X  element. 

In  the  corresponding  female  maturation  stages  a  tetrad  is  formed,  because 
there  are  two  of  these  chromosomes  present.  They  behave  normally  in 
maturation,  and  every  ripe  egg  receives  one  of  them.  Therefore,  after  fer- 
tilization there  will  be  two  types  of  embryos;  (i)  those  receiving  two  X 
chromosomes,  one  of  which  came  from  each  parent.  These  develop  into 
females.  (2)  Those  receiving  one  X  chromosome  from  the  mother,  and 
none  from  the  father.  These  are  the  males. 

"Sex-linked"  Inheritance. — This  is  a  peculiar  and  instructive  type  of 
inheritance,  which  behaves  in  a  way  suggesting  that  the  trait  may  be  carried 
in  association  with  the  X  chromosome.  Its  chief  distinction  is  that  it  is  never 
carried  from  father  to  son,  the  son  never  receiving  an  X  chromosome  from 
the  father.  The  best  human  example  is  hereditary  color-bindness,  which  is 
a  "sex-linked"  recessive  character,  due  to  some  lack  in  the  X  chromosome. 
The  presence  of  one  normal  color  bearing  X  chromosome  is  sufficient  to 
establish  color  vision. 

Changes  Following  Impregnation. — The  process  of  impregnation 
of  the  ovum  has  been  observed  most  accurately  in  the  lower  types. 
The  process  is  as  follows:  The  head  of  a  single  spermatozoon  joins 
with  an  elevation  of  the  yolk  substance,  the  tail  remaining  motionless  and 


CHANGES   FOLLOWING   IMPREGNATION 


787 


then  disappearing.  The  head  enveloped  in 
the  protoplasm  then  sinks  into  the  yolk  and 
becomes  a  nucleus,  from  which  the  yolk 
substance  is  arranged  in  radiating  lines. 
This  is  the  male  pronucleus.  The  middle 
piece  of  the  sperm  is  believed  to  furnish 
a  new  centrosome  to  the  ovum,  thus  re- 
storing its  capacity  for  cell  division.  The 
centrosome  now  divides  and  moves  to  either 
side  the  two  pronuclei,  and  a  segmenta- 
tion spindle  is  formed,  which  bears  all 
the  chromosomes  from  both  pronuclei. 
The  first  segmentation  occurs,  and  divides 
the  egg  into  two  cells,  hi  each  of  which  there 
is  the  unreduced  chromosomal  number. 

The  process  of  segmentation  begins 
almost  immediately  in  each  half  of  the 
divided  egg,  and  cuts  it  also  in  two.  The 
process  is  repeated  until  at  last  by  continued 
cleavages  the  whole  yolk  is  changed  into  a 
mulberry-like  mass,  still  enclosed  by  the 
zona  pellucida,  figure  502.  Fertilization 
probably  takes  place  in  the  Fallopian  tubes, 
and  segmentation  of  the  fertilized  ovum 
occurs  on  its  passage  to  the  uterus. 

The  passage  of  the  ovum  from  the  ovary 
to  the  uterus  occupies  probably  eight  or  ten 
days  in  the  human. 

The  peripheral  cells,  which  are  formed 
first,  arrange  themselves  at  the  surface  of  the 
yolk  into  a  membrane,  the  ectoderm.  The 
deeper  cells  of  the  interior  pass  gradually  to- 
ward the  surface,  thus  increasing  the  thick- 
ness of  the  membrane  already  formed  by  a 
second,  or  entoderm,  layer  of  cells,  while  the 
central  part  of  the  yolk,  the  blastoderm  cavitv, 
remains  filled  only  with  a  clear  fluid.  By 
this  means  the  yolk  is  shortly  converted  into 
a  kind  of  secondary  vesicle,  the  walls  of 
which  are  composed  externally  of  the  origi- 
nal vitelline  membrane,  and  within  by  the 
two  newly  formed  cellular  layers,  the  blasto- 
derm or  germinal  membrane,  as  they  are 


FIG.  502. — Conversion  of  the 
Morula  to  the  Blastula,  For- 
mation of  Blastodermic  Vesicle  and 
Membrane.  A,  Appearance  of 
segmentation  cavity  and  attach- 
ment of  inner  cell  mass  to  ectoderm 
at  upper  pole  of  ovum;  B1,  exten- 
sion and  flattening  of  inner  cell 
mass  as  it  occurs  in  rabbits  and 
some  other  mammals;  B2,  exten- 
sion of  entoderm  as  it  occurs  in 
insectivora,  monkeys,  apes,  and 
man;  C,  completion  of  bilaminar 
blastodermic  vesicle;  BC,  blasto- 
dermic  cavity;  EC,  ectoderm;  EE, 
embryonic  ectoderm;  EX,  ento- 
derm; 7,  inner  cell  mass;  SC,  seg- 
mentation cavity;  ZP,  zona  pellu- 
cida, (Cunningham.) 


788  DEVELOPMENT 

called.  A  third  cellular  layer,  the  mesoderm,  is  soon  developed  between 
the  other  two.  The  fetus  results  from  the  harmonious  growth  of  these  three 
layers,  each  of  which  is  the  source  of  certain  tissues  and  organs. 

Important  changes  occur  in  the  structure  of  the  mucous  membrane  of 
the  uterus.  The  epithelium  and  subepithelial  connective  tissue,  together 
with  the  tubular  glands,  increase  rapidly,  and  there  is  a  greatly  increased 
vascularity  of  the  whole  mucous  membrane,  while  a  substance  composed 
chiefly  of  nucleated  cells  fills  up  the  interf  ollicular  spaces  in  which  the  blood 
vessels  are  contained.  The  effect  of  these  changes  is  an  increased  thickness, 
softness,  and  vascularity  of  the  mucous  membrane,  the  superficial  part  of 
which  itself  forms  the  membrana  decidua. 

The  object  of  this  increased  development  is  the  production  of  nutritive 
materials  for  the  ovum;  for  the  cavity  of  the  uterus  shortly  becomes  filled 


FIG.  503. — Section  of  the  Lining  Membrane  of  a  Human  Uterus  at  the  Period  of 
Commencing  Pregnancy,  Showing  the  Arrangement  and  Other  Peculiarities  of  the  Glands, 
d,  d,  d,  with  Their  Orifices,  a,  a,  a,  on  the  Internal  Surface  of  the  Organ.  Twice  the 
natural  size. 


with  secreted  fluid,  consisting  almost  entirely  of  nucleated  cells  in  which  the 
chorion  villi  are  embedded. 

When  the  ovum  first  enters  the  uterus  it  becomes  embedded  in  the  structure 
of  the  decidua,  which  is  yet  quite  soft,  and  in  which  soon  afterward  three 
portions  are  distinguishable.  These  have  been  named  the  decidua  vera,  the 
decidua  basalis,  and  the  decidua  capsularis. 

In  connection  with  these  villous  processes  of  the  chorion  there  are  de- 
veloped depressions  or  crypts  in  the  decidua  vera,  which  correspond  in  shape 
to  the  villi  they  are  to  lodge;  and  thus  the  chorionic  villi  become  more  or  less 
embedded  in  the  maternal  structures.  These  uterine  crypts,  it  is  important 
to  note,  are  not,  as  was  once  supposed,  merely  the  open  mouths  of  the 
uterine  follicles. 

The  Placenta. — During  these  changes  the  deeper  part  of  the  mucous 
membrane  of  the  uterus,  at  and  near  the  region  where  the  placenta  is  placed, 
becomes  hollowed  out  by  sinuses,  or  cavernous  spaces,  which  communicate 
on  the  one  hand  with  arteries  and  on  the  other  with  veins  of  the  uterus.  Into 


THE  PLACENTA 


789 


these  sinuses  the  villi  of  the  chorion  protrude,  pushing  the  thin  walls  of  the 
sinuses  before  them,  and  so  come  into  intimate  relation  with  the  blood  con- 
tained in  them.  There  is  no  direct  communication  between  the  blood  vessels  of 
the  mother  and  those  of  the  fetus;  but  the  layer  or  layers  of  membrane  inter- 
vening between  the  blood  of  the  one  and  of  the  other  offer  no  obstacle  to  a 
free  interchange  of  matters  between  them  by  diffusion  and  osmosis.  Thus 
the  villi  of  the  chorion,  containing  fetal  blood,  are  bathed  or  soaked  in  mater- 
nal blood  contained  in  the  uterine  sinuses. 

The  placenta,  therefore,  of  the  human  subject  is  composed  of  a  fetal 
part  and  a  maternal  part — the  term  placenta  properly  including  all  that 
entanglement  of  fetal  villi  and  maternal  sinuses,  by  means  of  which  the 


Decidua  basal  is 
Unchanged  layer  Maternal  vessel 


Primitive  streak 
Mesoderm 

Placental  rillus 


Ectoderm 


Stratum  spongiosnm 
Stratum  compactum 

Placental  villus 


Villas- 


Cavity  of 
blastoderm 


Cavity  whicl 
becomes  coeloin 


•Mesoderm 


Decidua  vera/ 


Entodenn 
idua  vera 


FIG    504. — Diagram  of  the  Early  Stage  of  Human  Embryo  in  Relation  to  the  Uterus 

(Cunningham.) 


blood  of  the  fetus  is  enriched  and  purified  after  the  fashion  necessary  for  the 
proper  growth  and  development  of  those  parts  which  it  is  designed  to  nourish. 
The  whole  of  this  structure  is  not,  as  might  be  imagined,  thrown  off 
immediately  after  birth.  The  greater  part,  indeed,  comes  away  at  that  time, 
as  the  after-birth;  and  the  separation  of  this  portion  takes  place  by  a  rending 
or  crushing  through  of  that  part  at  which  its  cohesion  is  least  strong,  namely, 
where  it  is  most  burrowed  and  undermined  by  the  cavernous  spaces  before 
referred  to.  In  this  way  it  is  cast  off  with  the  fetal  membrane.  The  remain- 
ing portion  is  either  gradually  absorbed,  or  thrown  off  in  the  uterine  dis- 


7QO 


DEVELOPMENT 


charges  which  occur  at  this  period.  A  new  mucous  membrane  is  of  course 
gradually  developed. 

Circulation  of  Blood  in  the  Fetus. — The  circulation  of  blood  in  the 
fetus  differs  considerably  from  that  of  the  adult. 

Returning  from  the  placenta  by  the  umbilical  vein  the  blood  is  first  con- 
veyed to  the  under  surface  of  the  liver,  where  the  stream  is  divided — a  part  of 
the  blood  passing  straight  on  to  the  inferior  vena  cava  through  a  venous  canal 
called  the  ductus  venosus,  while  the  remainder  passes  into  the  portal  vein  and 
reaches  the  inferior  vena  cava  only  after  circulating  through  the  liver.  It  is 
carried  by  the  vena  cava  to  the  right  auricle  of  the  heart,  into  which  cavity 


FIG.  505. — Diagrammatic  View  of  a  Vertical  Transverse  Section  of  the  Uterus  at  the 
Seventh  or  Eighth  Week  of  Pregnancy,  c,  c,  c',  Cavity  of  uterus,  which  becomes  the 
cavity  of  the  decidua,  opening  at  c,  c,  the  cornua,  into  the  Fallopian  tubes,  and  at  c'  into  the 
cavity  of  the  cervix,  which  is  closed  by  a  plug  of  mucus;  dv,  decidua  vera;  dr,  decidua 
reflexa,  with  the  sparser  villi  embedded  in  its  substance;  ds,  decidua  serotina,  involving 
the  more  developed  chorionic  villi  of  the  commencing  placenta.  The  fetus  is  seen  lying 
in  the  amniotic  sac.  The  umbilical  cord  and  its  vessels  pass  up  from  the  umbilicus  to 
the  distribution  of  the  blood  vessels  in  the  villi  of  the  chorion-  and  the  pedicle  of  the 
yolk-sac  the  cavity  between  the  amnion  and  chorion.  (Allen  Thomson.) 


the  blood  is  also  pouring  that  has  circulated  in  the  head  and  neck  and  arms, 
and  has  been  brought  to  the  auricle  by  the  superior  vena  cava.  It  might  be 
naturally  expected  that  the  two  streams  of  blood  would  be  mingled  in  the 
right  auricle,  but  such  is  the  case  only  to  a  slight  extent.  The  blood  from  the 


CIRCULATION  OF  BLOOD  IN  THE  FETUS  791 

superior  vena  cava — the  less  pure  fluid  of  the  two — passes  almost  exclusively 
into  the  right  ventricle,  through  the  auriculo- ventricular  opening,  just  as  it 
does  in  the  adult.  The  blood  of  the  inferior  vena  cava  is  directed  by  a  fold 
of  the  lining  membrane  of  the  heart,  called  the  Eustachian  valve,  through  the 
foramen  ovale  into  the  left  auricle  and  into  the  left  ventricle,  and  out  of  this 
into  the  aorta,  and  thence  to  all  the  body,  but  chiefly  to  the  head  and  neck. 
The  blood  of  the  right  ventricle  is  sent  out  in  small  amount  through  the  pul- 
monary artery  to  the  lungs,  and  thence  to  the  left  auricle,  as  in  the  adult, 


FIG.  506. — Diagram  of  the  Fetal  Circulation. 

but  the  greater  part  by  far  passes  through  a  canal,  the  ductus  arteriosus, 
leading  from  the  pulmonary  artery  into  the  aorta  just  below  the  origin  of  the 
three  great  vessels  which  supply  the  upper  parts  of  the  body,  and  is  distributed 


792 


DEVELOPMENT 


to  the  trunk  and  lower  parts  of  the  body.  A  large  portion  passes  out  by 
way  of  the  two  umbilical  arteries  to  the  placenta.  From  the  placenta  it  is 
returned  by  the  umbilical  vein  to  the  under  surface  of  the  liver,  from  which 
the  description  started. 

After  birth  the  foramen  ovale,  the  ductus  arteriosus,  and  ductus  venosus 
all  close,  and  the  umbilical  vessels  are  tied  off,  so  that  the  two  streams  of 
blood  which  arrive  at  the  right  auricle  by  the  superior  and  inferior  vena  cava, 
respectively,  thenceforth  mingle  in  this  cavity  of  the  heart,  and  pass  into 
the  right  ventricle,  by  way  of  the  pulmonary  artery  to  the  lungs,  and  through 


FIG.  507. — Dissection  of  the  Lower  Half  of  the  Female  Mamma  During  the  Period  of 
Lactation.  §. — In  the  left-hand  side  of  the  dissected  part  the  glandular  lobes  are  exposed 
and  partially  unravelled,  and  on  the  right-hand  side  the  glandular  substance  has  been 
removed  to  show  the  reticular  loculi  of  the  connective  tissue  in  which  the  glandular  lobules 
are  placed,  i,  Upper  part  of  the  mammilla  or  nipple;  a,  areola;  3,  subcutaneous  masses 
of  fat;  4,  reticular  loculi  of  the  connective  tissue  which  support  the  glandular  substance 
and  contain  the  fatty  masses;  5,  one  of  three  lactiferous  ducts  shown  passing  toward  the 
mammilla,  where  they  open;  6,  one  of  the  sinus  lartei  or  reservoirs;  7,  some  of  the  glandular 
lobules  which  have  been  unravelled;  7',  others  massed  together.  (Luschka.) 

these,  after  aeration,  to  the  left  auricle  and  ventricle,  to  be  distributed  over 
the  body. 

Parturition. — With  the  implantation  of  the  embryo  and  the  develop- 
ment of  the  placenta,  the  uterus  grows  rapidly  until  the  end  of  pregnancy. 
The  muscles  of  its  walls  increase  enormously  in  volume,  apparently  by 
an  increase  in  the  size  of  the  fibers,  and  the  whole  structure  may  become 
thirty  or  forty  times  its  size  in  the  resting  period.  Many  changes  take 
place  also  in  other  parts  of  the  body,  changes  which  are  dependent  on  the 


LACTATION 


793 


presence  of  the  fetus  and  a  change  in  the  number  or  quantity  of  hormones 
developed  during  gestation.  Full-term  pregnancy  occurs  when  the  uterus  is 
isolated  from  the  nervous  system,  hence  it  has  been  inferred  that  there  is 
some  sort  of  special  secretion  or  hormone  possibly  of  the  embryo  itself,  that 
makes  its  way  into  the  blood  and  influences  the  organs  of  the  mother. 

At  the  end  of  the  period  of  pregnancy  the  strong  uterine  walls  begin 
periodic  contractions  which  ultimately  result  in  the  delivery  of  the  fetus. 
These  contractions  are  at  first  weak  and  at  long  intervals,  but  gradually 
become  very  strong  and  even  violent  and  follow  each  other  in  rapid  succes- 
sion. The  uterine  contractions  are  supported  by  reflex  contractions  of  the 
abdominal  and  thoracic  muscles.  After  the  fetus  is  delivered  the  uterine 
contractions  become  milder,  but  still  continue  until  the  placenta  is  finally 
expelled. 


FIG.  508. 


FIG.  509. 


FIG.  508. — Section  of  Mammary  Gland  of  Bitch,  Showing  Acini,  Lined  with  Epithelial 
Cells  of  a  Polyhedral  or  Short  Columnar  Form.     X  200.     (V.  D.  Harris.) 
FIG.  509. — Globules  and  Molecules  of  Cow's  Milk.     X  400. 

The  initiation  of  the  contractions  of  the  uterus  at  delivery  probably  de- 
pends on  the  chemical  stimulation  of  some  substance  or  substances  produced 
in  the  uterus  itself  or  in  the  fetus;  substances  that  react  on  the  nervous  mech- 
anism and  on  the  uterine  muscles  themselves.  This  view  cannot  be  said  to 
be  proven,  but  it  is  supported  by  certain  observed  facts  and  experiments. 

Lactation. — There  is  a  marked  development  of  the  mammary  glands, 
during  the  period  of  gestation,  especially  in  the  later  weeks.  No  milk  is 
secreted  during  this  time  and  the  change  is  sharply  limited  to  the  processes  of 
growth  in  the  glands.  Upon  delivery  of  the  fetus  the  gland  immediately 
enlarges  very  sharply  and  an  abundant  secretion  is  formed.  During  gesta- 
tion it  is  thought  that  an  inhibitive  hormone  prevents  actual  secretion. 
After  parturition  these  hormones  are  removed  and  secretion  begins  under  the 
stimulative  influence  of  definite  hormones  from  the  corpus  luteum,  the  invo- 
luting uterus,  and  the  mammary  gland  tissue,  see  Chapter  on  ductless  glands. 


794  DEVELOPMENT 

The  secretion  of  the  first  few  days  is  called  the  colostrum.  It  contains  a 
larger  per  cent,  of  solids,  has  the  large  granular  colostral  corpuscles,  is  more 
alkaline  than  ordinary  milk,  and  has  a  specific  gravity  of  1040  to  1046. 

The  mammary  glands  have  been  isolated  from  the  nervous  system  to 
determine  whether  or  not  the  association  in  time  between  their  changes  and 
the  changes  in  the  uterus  were  of  a  nervous  nature.  The  isolated  mammae 
develop  and  begin  lactation  at  parturition  as  in  the  normal.  It  would  seem 
that  there  is  some  special  form  of  stimulation  through  the  medium  of  the 
blood,  i.e.,  by  the  hormones  as  suggested  in  the  preceding  paragraph.  Yet 
one  must  not  draw  the  conclusion  that  the  nervous  system  exerts  no  influence 
on  the  mammary  gland.  Stimulation  of  the  nerves  to  the  gland  produces 
vascular  changes  that  increase  or  decrease  the  quantity  of  milk  secreted. 
Many  observations  have  been  noted  in  women,  which  show  that  the  secretion 
of  milk  is  sharply  influenced,  or  even  completely  suppressed  by  nervous  states. 

The  Composition  of  Milk. — Milk  has  a  specific  gravity  of  1028  to 
1034.  Its  fat  is  held  in  emulsion.  Under  the  microscope,  it  is  found  that 
the  milk  globules  vary  in  size,  the  majority  being  from  2  to  3^  in  diameter. 
The  old  view  that  they  have  an  investing  membrane  of  albuminous  material 
is  now  generally  discarded. 

COMPOSITION  OF  COLOSTRUM  (PFEIFFER). 

Proteins 5.71 

Fat 2  . 04 

Sugar 3.74 

Salts 0.28 

Water 88.23 

100.00 

SALTS  IN  WOMAN'S  MILK  (ROTCH). 

Calcium  phosphate 23  . 87 

Calcium  silicate 1.27 

Calcium  sulphate 2.25 

Calcium  carbonate 2.85 

Magnesium  carbonate 3-77 

Potassium  carbonate 23  .47 

Potassium  sulphate 8.33 

Potassium  chloride 12.05 

Sodium  chloride 21.77 

Iron  oxide  and  alumina o-37 

100.00 

In  addition  to  the  oil  or  butter  fat,  milk  contains  certain  proteins,  milk- 
sugar,  and  several  salts.  Its  percentage  composition  is  given  in  the  tables 
appended. 


THE    COMPOSITION    OF    MILK  795 

CHEMICAL  COMPOSITION  OF  MILK.     (AFTER  FOSTER,  HARRINGTON,  et  al.) 

Human.       Cow.  Mare.  Bitch. 

Water 87.30         87  90  76 

Solids 12.70         13  10  24 

Fats 4.00           4.0  2.0         10.  o 

Proteins 1.50           4.0  2.5          10.0 

Sugar 7-°°           4-3  5-°           3-5 

Salts 0.20           0.7  0.5           0.5 


FAHRENHEIT 

and 

CENTIGRADE 
SCALES. 


F. 

500° 
401 


374 
356 
347 


811 
302 
284 
275 


230 
212 
203 
194 
176 
167 
140 
122 
113 
105 
104 
100 


98.5 

95 

86 

77 

68 

50 

41 


C. 

260° 
205 
200 
195 
190 
180 
175 
170 
365 
160 
155 
150 
140 
135 
130 
120 
115 
110 
100 
95 


50 

45 

40.54 

40 

37.8 


25 
20 
10 
5 
0 

-  5 

-  10 
-15 
-20 
-25 
-30 
-40 
-60 


1    deg.F.  =  .54°C. 
1.8      "    =    1°C. 
3.6       "    =    2°0. 
4.5       "    •-=    2.5°C 
5.4       "    =    3°C. 


To  convert  de- 
grees F.  into  de- 
grees C.,  subtract 
32,  and  multiply 
by|. 


To  convert  de- 
grees C.  into  de- 
grees F.,  multiply 
by  |,  and  add  32°. 


MEASUREMENTS. 

FEENCH  INTO  ENGLISH. 


LENGTH. 

1  metre 

10  decimetres    I    =  39.37  English 
100  centimetres  f  inches. 

1,000  millimetres  J  (or  1  yd.  and  3^  in.) 


1  decimetre     ) 

10  centimetres  V  =  3.937  inches 
100  millimetres  )  (or  nearly  4  inches.) 


1  centimetre    )    =  .3937  or  about 
10  millimetres   f       (nearly  g  inch.) 
1  millimetre         =  nearly  &  inch. 

OR, 

ONE  METRE  =  39.37079  inches. 
(It  is  the  ten-millionth  part  of  a  quarter 

of  the  meridian  of  the  earth.) 

1  Decimetre    =  4  in. 

1  Centimetre  =  j%  in. 

1  Millimetre    =  <&  in. 

Decametre      =  32.80  feet. 

Hectometre    —  109.36  yds. 

Kilometre       =  0.62  miles. 
One  inch  =  2.539  Centimetres. 
One  foot  =  3.047  Decimetres. 
One  yard  =  0.91  of  a  Metre. 
One  mile  =  1.60  Kilometre. 
The  cubic  centimetre  (1 5.432  grains— 1 
gramme)  is  a  standard  at  4°  C.,  the 
grain  at  16°.66  C. 


WEIGHT. 

(One  gramme  is  the  weight  of  a  cubic 
centimetre  of  water  at  4°  C.  at  Paris). 
1  gramme  ] 

10  decigrammes    i  =  15.432349  grs. 
100  centigrammes  (      (or  nearly  15^). 
1,000  milligrammes  J 


1  decigramme 
10  centigrammes  V  =  rather  more 
100  milligrammes  )     than  l^j  grain. 


1  centigramme    \  =  rather  more 
10  decigrammes    (     than  5%  grain. 


1  milligramme        =  rather  more 

than  3g,j  grain. 
OR 

1  Decigramme  =  2  dr.  34  gr. 

1  Hectogrm.      =  8^3  oz.  (Avoir.) 

1  Kilogrm.         =  2  Ib.  3  oz.  2  dr.  (Avoir.) 


A  grain  equals  about  1.16  gram., 

a  Troy  oz.  about  31  gram., 

a  Ib.  Avoirdupois  about  ^  Kilogrm., 

and  1  cwt.  about  50  Kilogrms. 


CAPACITY. 

1,000  cubic  decimetres  I  =  1  cubic 
1 ,000,000  cubic  centimetres  f      metre. 


1  cubic  decimetre      ) 

or  5-  =  1  litre. 

1,000  cubic  centimetres    } 

OR 

ONE  LITRE  =  1  pt.  15  oz.  1  dr.  40. 
(For  simplicity,  Litre  is  used  to  signify 
1  cubic  decimetre,  a  little  less  than  1 
English  quart.) 

Decilitre  (100  c.c.)          =  fy$  oz. 
Centilitre  (10  c.c.)  =  2f  dr. 

Millilitre  (1  c.c.)  =  17  m. 

Decalitre  =  2£  gal. 

Hectolitre  =  22  gals. 

Kilolitre  (cubic  metre)  =  27^  bushels. 
A  cubic  inch  =  16.38  c.c.  ;  a  cubic  foot 
=  28.315  cubic  dec.,  and  a  gallon  = 
4.54  litres. 


CONVERSION  SCALE. 

To  convert  GRAMMES  to  OUNCES  avoir- 
dupois, multiply  by  20  and  divide  by  567. 

To  convert  KILOGRAMMES  to  POUNDS, 
multiply  by  1,000  and  divide  by  454. 

To  convert  LITRES  to  GALLONS,  nul- 
tiply  by  22  and  divide  by  100. 

To  convert  LITRES  to  PINTS,  nit  Itiply 
by  88  and  divide  by  50. 

To  convert  MILLIMETRES  to  INCHES, 
multiply  by  10  and  divide  by  254. 

To  convert  METRES  to  YARDS,  multi- 
ply by  70  and  divide  by  64. 


SURFACE   MEASURE. 

1  square  metre  =  about  1550  sq.  inches. 
Or  10.000  sq.  centimetres,  or  10.75  sq.  ft. 
1  sq.  inch  =  about  6.4  sq.  centimetres. 
1  sq.  foot  =       "     930      " 


ENERGY  MEASURE. 

1  kilogrammetre=about  7.24  ft.  pounds 
1  foot  pound        =     "      .1381  kgin. 
1  foot  ton  =    "      310  kgm. 


HEAT  EQUIVALENT. 

1  kilocalorie  =  424  kilogrammetres. 


ENGLISH 

Apothecaries  Weight. 

7000  grains  =  1  Ib. 

Or 
437.5  grains  =  1  oz. 


MEASURES. 

Avoirdupois  Weight. 

16  drams      =  1  oz. 
16  oz.  =  1  Ib. 

28  Ibs.  =.  1  quarter. 

4  quarters  =  1  cwt. 
20  cwt.          =  1  ton. 


Measure  of  1  decimetre,  or  10  centimetres,  or  100  millimetres. 


1234  567 

The  micron  (symbol,  M)  IB  the  unit  of  microscopic  measurement  = 


796 


INDEX 


Abdominal  viscera,  vascular  nerves  for, 

257 

Abducens  nerve,  66 1 
Aberration,  chromatic,  760 

spherical,  760 
Absorption,  409 

coefficient  of,  304 

conditions  for,  409 

methods  of,  409 

of  carbohydrates,  408 

of  fats,  416 

of  minerals,  417 

of  protein,  414 

of  water,  417 

places  for,  409 

rapidity  of,  408 

through  the  intestines,  412,  414 
the  lungs,  418 
the  mouth,  410 
the  skin,  418 
the  stomach,  410 

Accelerator  centers  for  heart,  210,  620 
Accessory  olives,  615 

thyroids,  484 

Accommodation  of  vision,  728,  730 
Achromatic  layer,  18 

spindle,  20 
Achro  matin,  18 
Achroodextrin,  356 
Acromegaly,  498 
Activating  ferments,  347 
Adamkiewicz  reaction,  107 
Adenase,  94 
Adenine,  93 
Adenoid  tissue,  36 
Adipose  tissue,  37,  38 
Adrenalin,  490 
Adrenals,  488 

Afferent  fibers  in  the  sympathetics,  672 
After-birth,  788 

-images,  741,  748,  765 

-sensations,  686 
Agglutinative  substances,  151 
Air,  284 

analysis,  297 


Air,  changes  in,  during  respiration,  297, 298 

complemental,  294,  328 

composition  of,  297 

diffusion  of,  301 

pressure  of,  301,  325,  328 

quantity  breathed,  294,  328 

reserve,  295,  328 

residual,  295 

supplemental,  295 

tidal,  294,  328 

volume  breathed,  328 
Alanine,  81 
Albumin,  egg,  105 

properties  of,  no 

serum,  88 

in  urine,  435,  453 
Albuminate,  acid,  112 
Albuminoids,  87,  89 

tests  for,  no 
Albumins,  86,  88 

reactions  of,  1 10 
Albumoses,  96 
Alcohol  as  a  food,  345 
Alexin,  150 

Alkali  metaprotein,  112 
Amboceptor,  150 
Ameba,  3 
Ameboid  leucocytes,  133 

movement,  3,  4,  134 
Amino-acids,  81 
Amitosis,  19 
Ammonia,  effect  of  breathing,  318 

in  expired  air,  301 

in  the  urine,  433 
Ammonium  carbamate,  431,  462 
Amylolytic  ferments,  347 
Amylopsin,  347,  382 
Amylose,  101 
Anabolism,  7,  454 
Anabolites,  454 
Anacrotic  limb,  239 

wave,  239 
Anaphase,  21 
Anaphylaxis,  152 
Anelectrotonus,  536 


797 


798 


INDEX 


Animal  heat,  501 

starch,  102 

Animals  differentiated  from  plants,  12 
Anode,  537  ^ 

Ano-spinal  center,  597 
Anterior  association  center,  650 
Antiperistalsis,  395 
Antiscorbutic  substance,  481,  482 
Antisubstances  in  blood,  152 
Antithrombin,  124 
Antitoxins  in  blood,  151 
Apnea,  316,  333 
Arborization,  interepithelial,  74 
Archoplasm,  19 
Areolar  tissue,  35 
Arginine,  82 

Aristotle's  experiment,  756 
Arrhythmia,  sinus,  213 
Arterial  blood,  analysis  of,  134 
flow,  229 

rhythmic,  230 
velocity  of,  231 
gases  of,  301 
pulse,  272 

blood-pressure  and  nervous  regu- 
lation in,  274 
tone,  244 
Arteries,  173 

blood-pressure  in,  218,  269,  274 
coats,  173 
nerves  of,  1 74 
tone  of,  244 
Arterioles,  173 
Articulate  sounds,  555 
Aspartic  acid,  82 
Asphyxia,  316,  333 
Assimilation,  6,  7 

Association  centers  of  cerebral  cortex,  648 
Aster,  20 

Astigmatism,  736,  762 
Atmosphere,  composition  of,  297 
Atmospheric  pressure,  301 
Attraction  sphere,  19 
Auditory  center,  647 
judgments,  712 
nerve,  659 

Auricles,  action  of,  179 
Autolytic  substances,  150 
Autonomic  system,  663,  669 
Axis  cylinder,  67 
Axone,  583 

Bacteria  in  digestive  tract,  391 


Basal  metabolism,  476 
Basement  membrane,  23,  336 
Basket  cells,  619 
Basophile,  133 
Beri-beri,  480 
von  Bezold's  ganglia,  201 
Bidder's  ganglia,  201 
Bile,  386,  408 

acids,  386,  408 

action  of,  on  peptic  digestion,  404 

chemical  composition  of,  386 

coloring  matter  of,  386 

functions  of,  387 

mode  of  discharge  of,  388 
of  secretion  of,  388 

pigments  of,  387 

salts,  386 

secretion  of,  388 
Bilirubin,  387 
Biliverdin,  387 
Binaural  sensations,  712 
Binocular  vision,  751 
Bioplasm,  2 
Biuret  reaction,  108 
Bladder,  urinary,  427 
Blastema,  2 
Blastoderm,  10 
Blind  spot,  731,  763 
Blood,  117 

agglutinative  substances  in,  150 

analysis  of,  man,  135 

anaphylaxis,  152 

antisubstances  in,  152 

antitoxins  in,  151 

arterial  flow,  229 

buffy  coat,  120 

capillary  flow,  231 

carbon  dioxide  of,  307 

chemical  composition  of,  134 

circulation  of,  166,  208 
experiments  on,  260 
in  fetus,  790 

coagulation  of,  118,  161 
calcium  in,  122 
conditions  affecting,  124,  162 
fibrin  in,  119 
rapidity  of,  119,  124,  162 
theories  of,  121 

composition  of,  table,  134,  135 
tissue  extracts  on,  163 

corpuscles  of,  117,  126,  131,  164 
chemical  composition  of,  138 


INDEX 


799 


Blood,  corpuscles  of,  colorless,  131 

ameboid  movement  of,  133,  222 

chemical  composition  of,  138 

number  of,  126 

phagocytosis,  158 

varieties  of,  131 
enumeration  of,  158 
percentage  of,  160 
red,  126 

characters  of,  126 

chemical  composition  of,  139 

corpuscles    of,    red,    action    of 
reagents  on,  157 

development  of,  129 

number  of,  126 

origin  of,  129 

rouleaux  formation,  128 
defibrinated,  120 
differences      between     arterial     and 

venous,  148 
elimination    of    carbon    dioxide    by, 

307 

examination  of,  157 
ferments  in,  137 
flow,  arterial,  229 

capillary,  231 

regulation  of,  242 

velocity  of,  235 
in  arteries,  231 
in  capillaries,  234 
in  veins,  235 

venous,  234 
gas  apparatus,  303 
gases  of,  301 
hemoglobin,  139 
human  types,  150 
isotonicity  of,  161 
laboratory  experiments  on,  157 
microscopical  examination  of,  157 
morphology  of,  126 
opsonins  in,  151 
oxygen  of,  307 
phagocytes,  135 
plasma,  117 

chemistry  of,  136,  163 

composition  of,  136 

percentage  of,  160 

physical  factors  of,  152 

reaction  of,  161 
platelets,  135 
portal,  148 
precipitins  in,  151 


Blood,  pressure,  218 

apparatus,  226 

arterial,  218,  269,  271,  272 

capillary,  227,  273 

in  man,  226,  227 

model,  271 

respiratory  undulations  of,  222 

variations  in,  228 

venous,  227 

production  of  heat  by  the,  503 
quantity  of,  117 

influence  on  secretion,  339 
respiratory  changes  in,  289 
serum,  118,  138,  152 

chemistry  of,  163 

composition  of,  135 

globulicidal  properties  of,  149 
specific  gravity  of,  161 
thrombocytes,  135 
transfusion,  149 
uses  of,  117 

variations  in  composition  of,  148 
velocity  of  flow,  235 
venous  flow,  234 
whipped,  162 
Blushing,  247 

Body,  chemical  composition  of,  79 
energy  requirements  of,  476 
experiments  on  the  chemistry  of,  105 
Bone,  42 

blood-vessels  of,  43 
canaliculi  of,  44 
cells,  45 
compact,  50 
development  of,  46 
growth  of,  51 
Haversian  canals  of,  44 
lacunae  of,  44,  45 
lamellae  of,  44,  45 
marrow,  42 

microscopic  structure  of,  44 
ossification  in  cartPage,  47 

in  membrane,  46 
periosteum  of,  43 
structure  of,  44 
Bowman's  sarcous  elements,  61 

theory  of  urinary  secretion,  436 
Brain,  603,  605,  613 

arrangement  of  different  parts,  603 
association  centers  of,  648 
distinctive     characters     of    human, 
605,  641 


8oo 


INDEX 


Brain,  fore-,  605,  625 

hind-,  605 

mid-,  606,  623 

motor  areas  of,  641 
centers,  641 
tracts  in,  621 

Rolandic  area  of,  644 

sensory  areas  of,  644 

vascular  nerve-supply  of,  253 

weight  of,  632 
Bread,  composition  of,  344 
Bronchi,  280 
Buffy  coat,  120 

Bulb,  the,  611;  and  see  Medulla. 
Bundle  of  His,  203 
Burdach,  tract  of,  587 

Caffeine,  93 
Cajal,  cells  of,  629 
Calcification,  47,  51 
Calcium  salts  in  the  body,  103 

in  coagulation  of  the  blood,   124 

tests  for,  115 
Calorimeter,  477 
Canaliculi,  44 
Cane  sugar,  100 
Capillaries,  174 

blood-pressure  in,  227,  273 

structure  of,  175 
Capillary  circulation,  272 

flow,  231 

velocity  of,  234 
Carbohydrates,  98,  113 

absorption  of,  by  intestines,  417 

as  foods,  344 

chemical  reactions  of,  113 

energy  value  of,  467 

metabolism  of,  467 

relation  of  epinephrin  to,  491 
Carbon,  amount  excreted,  456 

dioxide,  determination  of,  329 
elimination  of,  309,  320 

monoxide,  effect  of  breathing,  318 
hemoglobin,  142,  165 

in  proteins,  106 
Carbonates  in  the  body,  104 
Cardiac  force  of  action,  195 

conducting  system,  203 

conduction  stimulus,  206 
rate  of,  207 

contractions,  automatic,  265 
experiments  on,  260 


Cardiac    force    of    action,    contractions, 

maximal,  199 
cycle,  179,  193 
fibrillation,  209 

glands,  of  stomach,  363 

impulse,  185 

muscle,  62,  195,  529,  568 
action  of,  529 

automatic  contractions  of,  265 
development  of,  65 
properties  of,  195 
refractory  phase,  52 

nerves,  210,  266 

orifice  of  stomach,  374 
Cardiac  nodes,  203 
Cardio-accelerator  centers,  213,  615 
Cardiogram,  187,  261 
Cardiograph,  185 

Cardio-inhibitory  centers,  213,  614 
Cartilage,  39 

development  of,  41 

elastic,  41 

embryonic  spongy  bone,  48 

hyaline,  39 

temporary,  41 

vascularization  of,  48 

white  fibro-,  41 
Catacrotic  limb,  239 

wave,  240 

Catelectrotonus,  536 
Cathode,  535,  538 
Cauda  equina,  580 
Caudate  nucleus,  627 
Cell,  i,  8,  17,  18 

body,  17 

connection,  modes  of,  23 

difference    between    plant    and    ani- 
mal, 12 

division  of,  10,  19 

functions  of,  u,  14 

growth,  7,  10 

mutiplication,  19 

nucleus  of,  18 

structure  of,  17,  18 
Cells,  decay  and  death  of,  23 

modes  of  connection,  23 

origin  of,  23 

shapes  of,  22 

types  of,  22 
Cellulose,  13,  103 
Center  for  muscle  tone,  596 
Centers,  motor,  639,  641 


INDEX 


80 1 


Centers,  sensory,  644 

spinal,  596 
Centrosome,  19 

Cerebellar  cortex,  paths  through,  619 
Cerebellum,  617 

connection  with  bulb,  611,  615 

functions  of,  620 

general  structure  of,  618 
Cerebral   cortex,   association   centers  of, 
648 

localization  of  motor  function  of,  639 
of  sensory  function  of,  644 

nerve  cells  in,  629 

structure  of,  627 
Cerebrum,  627 

connection  with  bulb,  611 

effects  of  removal  of,  635,  637 

functions  of,  635 

motor  areas  of  cortex,  639,  641 

peduncles  of,  623 

sensory  areas  of,  644 

weight  of,  632 
Cerumen,  444 
Ceruminous  glands,  444 
Chemical   composition   of  the   body,    79 

elements  in  the  body,  79 

structure  of  proteins,  So 
Chemistry  of  the  body,  experiments  on,  105 
Chest,    changes   in   diameter   of,    during 

respiration,  325,  327 
Cheyne-Stokes  breathing,  317 
Chlorides  in  the  body,  103 

in  the  urine,  435,  450 

tests  for,  115 

Chlorine,  effect  of  breathing,  318 
Chlorophyll,  13 
Cholesterol,  98,  114 
Choline,  98 

Chorda  tympani,  350,  399 
Chordae  tendineae,  173 
Chromatic  aberration,  735,  761 
Chromatin,  18 
Chromophanes,  745 
Chromophiles,  72,  73 
Chromoplasm,  18 
Chromoproteins,  92 
Chromosomes,  20,  785 

and  Mendelian  inheritance,  785 
Chyme,  374 
Cilia,  31 
Ciliary  apparatus,  717 

contraction,  532,  570 


Circulation,  coronary,  215 

atmospheric  pressure  on,  318 

during  sleep,  673 

effect  of  respiration  on,  323 

fetal,  790 

in  brain,  253 

in  erectile  structures,  258 

laboratory  experiments  on,  260 

of  blood,  1 66,  218 

regulation  of  blood  flow,  242 

through  blood-vessels,  218 

time  of,  270 

vegetable,  5 

velocity  of,  234 
Coagulated  proteins,  96 
Coagulating  ferments,  347 
Coagulation  of  blood,  118,  161 

calcium  salts  in,  122 

conditions  affecting,  124,  162 

rapidity  of,  119,  124,  162 

theories  of,  121 
Cochlea,  704 
Cohnheim's  areas,  62 
Cold,  influence  of  extreme,  505 
Collagen,  90,  no 
Collaterals,  69 
Colloidal  solution,  85 
Colloids,  85,  152 
Color,  after-images,  747,  765 

-blindness,  748,  763 

complemental,  748 

extent  of  visual  field  for,  747 

Hering's  theory  of,  750 

limits  of  field  of  vision  for,  747,  765 

-mixing,  766 

sensations  of,  748 

vision,  theories  of,  749 

Young's  and  Helmholtz's  theory,  749 
Colorless  corpuscles,  131 
Colostrum,  794 
Comma  tract,  587 
Common  sensations,  679 
Compact  bone  formation,  50 
Complement,  150 
Complemental  air,  294,  328 

colors,  747 

Conducting  system,  cardiac,  203 
Conductivity  of  muscle,  513 
Conjugated  proteins,  87,  92 
Conjunctiva,  716 
Connective  tissues,  33 

adenoid,  36 


SO  2 


INDEX 


Connective  tissues,  adipose,  37 

areolar,  35 

cells  of,  33 

fibrous,  37 

gelatinous,  36 

general  structure  of,  33 

intercellular  substance  of,  34 

lymphoid,  36 

retiform,  36 

varieties  of,  34 

white  fibrous,  34,  37 

yellow  elastic,  35,  37 
Consonants,  555 
Contractility  of  muscle,  512 
Contraction  phase  of  muscle,  514 
Contracture,  524 
Convoluted  tubule,  423 
Cooking,  effects  of,  345 
Cornea,  716 
Corneoscopy,  766 
Corona  radiata,  624 
Coronary  circulation,  215 
Corpora  cavernosa,  258 

geniculata,  626 

quadrigemina,  625 

striata,  627 
Corpus  Arantii,  173 

dentatum,  617 

luteum,  500,  779 

internal  secretion  of,  500 

spongiosum,  258 
Corpuscles,  blood,  117,  126,  131,  164 

of  Golgi,  77 

of  Krause,  76 

of  Meissner,  75 

of  Pacini,  75,  379 
Corti,  organ  of,  704 
Coughing,  center  for,  614 
Cranial  nerves,  650 
Crassamentum,  118 
Creatinin,  433,  459,  463 
Cretinism,  485 
Crura  cerebri,  623 
Crusta,  623 

petrosa,  53 

phlogistica,  120 
Crystalloids,  152 
Crystals,  protein,  105 
Cutis  vera,  441 

anserina,  58 
Cystin,  81 

in  urine,  435 


Cytolysis,  149 
Cytoplasm,  9,  17 
Cytosine,  93 

Death,  7 
Decay,  7 
Decidua  basalis  and  capsularis,  788 

menstrualis,  779 

vera,  788 

Decussation  of  the  pyramids,  607 
Defecation,  396 

center  for,  597 
Degeneration  in  spinal  cord,  586 

reaction  of,  539 

Wallerian,  575,  586 
Deglutition,  358 

center  for,  614 

nervous  mechanism  of,  360 

time  occupied  in,  359 
Demarcation  currents,  516 
Dendrites,  65,  577 
Dental  papilla,  56 
Dentine,  53 
Depressor  nerve,  248 
Development,  781 
Dextrin,  101 

tests  for,  113 
Dextrose,  99 

tests  for,  113 

in  urine,  435 
Diabetes  mellitus,  470 
Dialysis,  152 
Diapedesis,  234 
Diaphragm  in  respiration,  288 
Diastole  of  heart,  179,  181 
Dicrotic  notch,  239 

pulse,  242 
Diet,  normal,  requisites  of,  454,  474 

tables,  475 
Diffusion,  152 
Digestion,  341,  346 

enzymes  in,  346 

experiments  in,  399 

in  intestines,  376,  390,  406 

in  mouth,  347,  399 

in  stomach,  360,  370 
Digestive  enzymes,  347 
Diphasic  current,  517 
Diplopia,  653,  751 
Disaccharides,  99 
Disassimilation,  7 
Distance,  estimation  of,  755 


INDEX 


8o3 


Diuretics,  action  of,  439 
Dogiel's  cells,  202 
Dreams,  673 

Ductless    glands,    influence    on    metab- 
olism, 482 
Ductus  arteriosus,  790 

deferens,  769 

venosus,  790 
Dyspnea,  315,  333 

Ear,  cochlea  of,  704 

external,  708 
function  of,  708 

internal,  703 
function  of,  710 

membranous  labyrinth,  704 

middle,  708 

function  of,  708 

organ  of  Corti,  705 

ossicles  of,  701 

semicircular  canals,  704 

tympanum,  701 
Edestin,  105,  no 
Egg  albumin,  105 
Eggs,  composition  of,  343 
Eighth  cranial  nerve,  659 
Elasticity  of  muscle,  511 
Elastin,  90,  in 
Electrocardiogram,  normal,  205 

of  extraventricular  systole,  209 

of  fibrilation,  208 
Electrotonus,  536,  566 
Elements,  chemical,  in  body,  79 
Eleventh  cranial  nerve,  662 
Embryonic  spongy  bone  cartilage,  48 
Emulsification,  114,  382 
Enamel,  54 

cap,  57 

germ,  56 

organ,  56 

papilla,  56 
Encephalon,  605 
End-brushes,  70 

-bulbs,  76 

-plates,  63 

Endocardiac  pressure,  187,  190 
Endocardium,  168 
Endomysium,  60 
Endoneurium,  69 
Endothelium,  26 
Energy,  income  and  output  of,  474 

requirements  for  body,  476 


Enteric  system,  671 
Enterokinase,  347,  381,  390,  409 
Enzymes,  347 

activating,  347 

amylolytic,  347 

coagulating,  347 

digestive,  347 

lipolytic,  347 

of  pancreatic  juice,  383,  407 

proteolytic,  347 
Eosinophile,  116,  133 
Epiblast,  n 
Epidermis,  441 
Epiglottis,  280 
Epinephrin,  490 
Epineurium,  69 
Epithelial  tissues,  24 
Epithelium,  24 

ciliated,  30,  32 

classification  of,  24 

columnar,  25,  27,  29 

cubical,  24 

functions  of,  25,  32 

glandular,  30 

simple,  24,  25 

situations  of,  24,  31 

specialized,  30 

squamous,  24,  28 

stratified,  24,  27 

transitional,  29 
Equilibrium,  sense  of,  68 
Erection  center,  598 
Erepsin,  347,  390 
Ergograph,  525 
Erythroblasts,  130 
Erythrocytes,  126 
Erythrodextrin,  356 
Eustachian  tube,  701,  709 

valve,  169,  791 
Excreta,  analysis  of,  456 

channels  of  elimination  of,  456 
Excretion,  421 

during  starvation,  473 

from  skin,  444 

laboratory  experiments  in,  448 
Expiration,  290,  297 

forced,  297 

muscles  of,  force  of,  288 

relative  time  of,  293 
Expired  air,  carbon  dioxide  of,  298,  330 

oxygen  of,  298 
External  genitals,  vascular  nerves  for,  258 


8o4 


INDEX 


Eye,  715 

anatomy  of,  716 

astigmatism,  731,  736,  762 

chromatic  aberration  of,  761 

image  formation,  725 

movements  of,  733 

muscles  concerned  in,  733 

optical  apparatus,  724 
axis,  727 

refractive  media  and  surfaces,  724 

schematic,  727 

spherical  aberration  of,  761 
Eyeball,  716 

blood-vessels  of,  723 

ciliary  apparatus,  717 

cornea,  716 

iris  and  lens,  717 

retina,  719 
Eyelids,  716 

Facial  nerve,  657 

function  of,  657 
paralysis  of,  657 
relation  to  taste,  658 
secretory  fibers,  657 
Falsetto  voice,  554 
Far-point,  760 
Fasciculus  antero-lateralis,  587 

cerebello-spinalis,  587 

cuneatus  and  gracilis,  587,  606 

of  Rolando,  606 

solitarius,  659 
Fasting,  472 

Fatigue,  effect  on  muscular  contraction, 
561,  564 

of  nerve  fiber,  534 
Fats,  96,  114 

absorption  of,  by  intestines,  518 

as  food,  341,  344 

destination  of,  467 

digestion  of,  382,  387 

emulsification  of,  114,  382,  387 

energy  value  of,  464 

metabolism  of,  463 

mobilization  of,  in  body,  466 

saponification  of,  114,  382 

source  of,  in  body,  465 

tests  for,  114 
Fatty  acids,  tests  for,  114 
Feces,  393 

composition  of,  393 

excretion  by,  455 


Fermentation,  391 
Ferments  in  the  blood,  137 

unorganized,  346,  and  see  Enzymes. 
Fetus,  circulation  of  blood  in,  790 
Fibers  of  Remak,  67 
Fibrillation,  auricular,  209 
Fibrin,  120 

ferment,  121 
Fibrinogen,  121,  137 
Fictitious  feeding,  365 
Fifth  cranial  nerve,  653 
Fillet   616 
Filtration,  411 
Filum  terminale,  55 
Finger,  vasomotor  changes  in,  276 
Fish,  composition  of,  342 
Flutter,  auricular,  209 
Food,  and  digestion,  341 

effects  of  deprivation  of,  472 

mastication  of,  348 

salts  of,  344 

time  of  passage  through  alimentary 

canal,  397 
Foods,  341 

carbohydrates,  344 

classification  of,  341 

effect  of  cooking,  345 

fats,  344 

fuel  value  of,  342 

heat  production  from,  342,  479 

income  and  output  of  energy,  455 

inorganic,  344 

liquid,  345 

mineral,  344 

nitrogenous,  341 

percentage  composition  of,  342 

salts,  344 

water,  345 

Forced  movements,  623 
Fore-brain,  625 

Form,  estimation  of,  653,  783,  754 
Fossa  ovalis,  169 
Fourth  cranial  nerve,  653 
Fovea  centralis,  719 
Frontal  association  center,  649 
Fructose,  100 

Galactose,  100 
Gall-bladder,  386 

emptying  of,  388 
Gallstone,  389 
Ganglia,  578 


INDEX 


8oS 


Ganglia,  Bezold's,  201 
Bidder's,  201 
Remak's,  201 
spinal,  functions  of,  595 
Ganglion  cells,  73 
Gases  in  alimentary  canal,  394 

in  blood,  determination  of,  334 
Gastric  digestion,  360,  402 

changes  in  food  in,  373 

circumstances  influencing,  372 

cleavage  products  of,  404 

products  of,  371 

time  of,  373 
hormone,  3.66 
juice,  368,  402 

acid  of,  368,  370 

action  on  milk,  372 

on  proteins,  370,  3  71,  404 

artificial,  404 

chemical  composition  of,  368,  403 

digestive  action  of,  402 

enzyme  action  of,  372,  404 

fictitious  meals,  action  on,  365 

hydrochloric  acid  in,  370 

pepsin  in,  372 

psychic  secretion  of,  403 

quantity  of,  368 

reaction  of,  368 

secretion  of,  365,  402 

quantity,  368 
lipase,  action  of,  372 
secretion,  changes  in  glands  during, 

365 

nervous  mechanism  of,  366 
Gelatin,  in 

effect  of  diet  of,  460 
Gelatinous  tissue,  36 
Genito-spinal  center,  598 
Germinal  membrane,  787 

matter,  2 

spot,  775 

vesicle,  773 

Gestation,  relation  of  to  internal  secre- 
tions, 500 
Giant  cells,  42 
Glands,  cardiac,  363 

gastric,  364 

mammary,  793 

pyloric,  364 

reproductive,    relation     to     metabo- 
lism, 498 

salivary,  348 


Glands,  sebaceous,  444 

secreting,  336 

sudoriferous,  443 

types  of,  336 
Gliadin,  83 
Globin,  145 
Globulins,  86,  89,  no 
Glomerulus,  422 
Glosso-pharyngeal  nerve,  659 

in  respiration,  312 
Glottis,  544 

respiratory  movements  of,  291 
Glucose,  99 
Glutaminic  acid,  82 
Glutelins,  86 
Glutenin,  83 
Glycemia,  470 
Glycocoll,  81,  386 
Glycogen,  468 

destination  of,  470 

formation  of,  468 

from  carbohydrate,  462 

from  protein,  459 

relation  to  metabolism,  469 

sources  of,  468,  469 

tests  for,  113 
Glycogenesis,  468 

hormone  control,  468 
Glycoproteins,  87,  95,  in 
Glycosuria,  469,  470 
Goblet  cells,  27 
Golgi,  corpuscles  of,  77 
Goll,  tract  of,  587 
Cowers'  tract,  587 
Graafian  follicles,  774 
Grape  sugar,  99 
Guanase,  94 
Guanilic  acid,  93 
Guanine,  94 

Haversian  canals,  44 

Head,  vascular  nerve  supply  of,  253 

Hearing,  699 

acuteness  of,  758 

limits  of,  708,  758 
Heart,  167 

accelerator  nerve  of,  213,  615 

action  of,  179,  195 

anatomy  of,  167 

automaticity  of,  204,  263,  265 
rate  of,  217,  261 
sequence,  261 


8o6 


INDEX 


Heart,  automaticity  of,  theories  of,  200 
-block,  202 
capacity  of,  1 70 
chambers  of,  168 
character  of  contraction,  195,  199 
compensatory  pause,  200 
conducting  bundle,  203 
coronary  circulation  of,  215 
cycle  of,  179,  193 
depressor  nerve  of,  248 
development  of,  171 
endocardiac  pressure,  187 
excised,  experiments  on,  263,  264 
force  of  action,  195 
frequency  of  action,  179,  217 
ganglia  of,  201 
impulse  of,  185 

influence    of    accelerator    nerve    on, 
210,  615 

of  coronary  circulation  on,  215 

of  drugs  on,  217 

of  inhibitory  nerves  on,  210,  614 

of  mechanical  tension  on,  216 

of  nervous  system  on,  210,  266 

of  nutrient  fluids  on,  263 

of  sympathetic  system  on,  210 

of  temperature  on,  216 

of  vagus  on,  210 
inhibitory  nerve  of,  210,  614 
irritability  of,  199 
isolated,  200,  263,  264 
metabolism  of,  204 
muscle,  62 

properties  of,  195 
production  of  heat  by,  503 
relation  of  rhythm  to  nutrition,  204 
rhythmic  contraction  of,  196,  204 
size  of,  170 
sounds  of,  183 

causes  of,  184 
structure  of,  171 
tonicity  of,  198 
valves  of,  172 

action  of,  181 
volume  of,  264 
weight  of,  1 70 
work,  per  diem,  479 
Heat,  animal,  501 

dissipation  of,  503 

from  lungs,  506 

from  skin,  504 
influence  of  extreme,  505 


Heat,  of  nervous  system  on  production 
of,  5°7 

produced  in  muscular  contraction ,  501 

-producing  organs,  503 

production  of  body-,  503,  506 

regulation  of  body-,  503 
centers  for,  507 

-rigor,  513,  526 

variations  in,  506,  507 
in  loss  of,  503 
in  production  of,  506 

Heidenhain's  experiments  on  urine  secre- 
tion, 436 
Height  weight  chart,  479 

formula  for  area,  479 
Heller's  ring  test,  108 
Hemacytometer,  159 
Hematin,  146 
Hematoblasts,  128 
Hematoidin,  146 
Hematoporphyrin,  146 
Hemianopsia,  646 
Hemin,  146 
Hemiopia,  646 
Hemochromogen,  145 
Hemoglobin,  87,  95,  139,  160,  307 

action  on  gases,  142 

combining  power  with  oxygen,  305, 
306 

crystals,  140,  16.4 
preparation  of,  105 

derivatives  of,  145,  164 

estimation  of,  142,  160 

reduced,  142 
Hemoglobinometer,  142,  160 

Dare's,  144 

FleischPs,  142 

Talquist's,  144 
Hemolysis,  149 
Henle's  membrane,  174 

loop,  423 

Hepatolytic  sera,  150 
Heredity,  784 
Hind-brain,  605 
Hippuric  acid,  433 

formation  of,  433-463 
His,  bundle  of,  202 
Histidine,  82 
Histones,  87,  91 
Hopkins-Cole  reaction,  107 
Hormones,  483 

gastric,  366,  36; 


INDEX 


807 


Hormones,  pancreatic,  491 

thyroid,  486 
Hyaline  cartilage,  39 

cells,  133 

leucocytes,  133 
Hyaloplasm,  9,  17 
Hydrochloric  acid,  370,  371 

combined,  371 

digestive  action  of,  371 

test  for  free,  371 
Hydrogen,  amount  excreted,  455 

in  proteins,  106 
Hydrolytic  cleavage,  83 
Hypermetropia,  738 
Hyperpituitarism,  496,  497,  498 
Hyperpnea,  316,  317 
Hyperthyroidism,  485 
Hypertonic  solutions,  154 
Hypoblast,  n 
Hypoglossal  nerve,  663 
Hypopituitarism,  496 
Hypothyroidism,  485 
Hypotonic  solutions,  154 
Hypoxanthine,  94 

Immune  body,  149 

Impregnation,   changes  in  ovum  follow- 
ing, 786 

Indican  in  urine,  test  for,  452 
Indol,  408 
Induction  coil,  557 
Infundibulum,  284 
Inheritance,  sex-linked,  786 
Inhibition  of  reflex  actions,  598 
Inorganic  foods,  344 

substances,  103 
Insalivation,  348 
Inspiration,  287,  288 

forced,  288 

muscles  of,  288 

quiet,  288 

relative  time  of,  293 
Intercellular  substance,  23,  34 
Interepithelial  arborizations,  74 
Internal,  capsule,  624 

ear,  703 

secretions,  483 

Intestinal  digestion,  376,  390 
role  of  bile  in,  387 

gases,  394 

juices,  406,  408 

secretion,  390 


Intestines,  absorption  in,  416 

action  of  microorganisms  in,  391 

digestion  in,  376 

feces  in,  393 

fermentation  in,  391 

gases  in,  394 

large,  summary  of  digestive  changes 
in,  3Qi 

movements  of,  394 
influence   of   nervous   system    on, 
396 

putrefaction  in,  392 

small,  summary  of  digestive  changes 
in,  390 

vascular  nerves  for,  257 
Intonation,  552 
Inulin,  1 02 
Invertase,  347 

Inverted  image  on  retina,  760 
Involuntary  muscle,  529,  569 
Iodine,  104 
Iris,  717 

contraction  of,  733 
Iron,  104 

tests  for,  115 
Irritability,  5 

of  heart-muscle,  199 

of  muscle,  512,  519,  559 
Islands  of  Langerhans,  378,  379,  492 
Isotonic  solutions,  154 
Isotonicity  of  blood,  161 
Ivory,  53 

Judgment   of   form   and   size    of   bodies, 
689 

of  form  and  solidity,  754 

of  size  and  distance,  755 
Jumping,  543 

Karyokinesis,  19 
Karyolymph,  18 
Karyoplasm,  18 
Karyosomes,  18 
Katabolism,  7,  454 
Katabolites,  454 
Keratin,  89,  no 
Kidneys,  421 

action  of  diuretics  en,  439 

blood  supply  of,  424 

effect  of  blood  pressure  on,  448 

factors  affecting  excretion  from,  436 

function  of,  421,  422 


8o8 


INDEX 


Kidneys,  internal  secretion  of,  493 

Mulpighian  bodies  of,  425,  426 

nerves  of,  426 

plethysmogram  of,  277 

structure  of,  421 

tubuli  uriniferi  of,  422 

vasa  efferentia  of,  426 
recta  of,  426 

vascular  nerves  of,  257 

volume  of  urine,  428 
Krause,  corpuscles  of,  76 

membrane  of,  61 

Kronecker-Meltzer     theory    of     degluti- 
tion, 358 
Kymograph,  220 

Labyrinth,  703 
Lachrymal  apparatus,  716 
Lactase,  347 
Lactation,  793 

relation  of  internal  secretion  to,  500 
Lacteals,  413 
Lactose,  101 
Lacunae,  44,  45 

Langerhans,  islands  of,  378,  379,  492 
Large    intestine,    summary    of    digestive 

changes  in,  391 
Laryngoscope,  549 
Larynx,  279,  544 
Latent  period  of  muscle,  514,  564 
Law,  DuBois,  566 

Pfluger's,  536,  566 
Leaping,  543 
Lecithins,  95,  97,  114 
Lecithoproteins,  87,  95 
Legumes,  composition  of,  344 
Lens  of  eye,  717 
Lenticular  nucleus,  627 
Leucin,  82 
Leucocytes,  116,  131 

number,  131 

phagocytic  action,  133 

varieties,  132 
Leucolytic  sera,  150 
Leucosin,  83 

Levers,  action  of,  in  the  body,  540 
Levulose,  100 
Lichinin,  102 

Liebermann's  reaction,  107 
Life,  phenomena  of,  i 
Limbs,  vascular  nerves  for,  258 
Linin,  18 


Lipase,  347,  382,  383 
Lipogenesis,  466,  493 
Lipoids,  98,  114 
Lipolytic  ferments,  347 
Liquid  foods,  345 
Liquor  sanguinis,  117 
Liver,  385 

glycogenic  function  of,  460 

internal  secretion  of,  493 

secretions  of,  384,  388 

structure  of,  385 

urea  formation  in,  461 

vascular  nerves  for,  257 
Localization,  cerebral,  639,  644 
Locomotion,  539 
Locus  ceruleus,  616 

Ludwig's  theory  of  urine  secretion,  436 
Lungs,  282 

absorption  from,  420 

blood  supply  of,  286 

excretion  by,  456 

interchange  of  gases  in,  309 

loss  of  heat  from,  506 

lymphatics  of,  286 

nerves  of,  287 

structure  of,  288 
Luxus  consumption,  458 
Lymph,  155 

chemical  composition  of,  155 

flow,  156 

formation  of,  155 
Lymphagogues,  156 
Lymphatic  sheaths,  perivascular,  179 

spaces,  in  blood-vessels,  1 79 
Lymphocyte,  116,  133 
Lymphocytes,  variety  of,  132 
Lymphoid  tissue,  36 
Lysine,  82 
Lysis,  149 
Lytic  substances,  149 

Magnesium  salts  in  the  body,  103,    115 

Malpighian  bodies,  426 

Maltase,  347,  355,  379,  382 

Maltose,  100,  356 

Malt  sugar,  100,  356 

Mammary  glands,  792,  793,  794 

Manometer,  192,  219 

Marrow,  bone,  42 

Mast  cells,  132 

Mastication,  348 

nervous  mechanism  of,  349 


INDEX 


8o9 


Mastoid  cells,  701 
Maximal  stimulus,  519 
Measures  and  weights,  table  of,  796 
Meat,  composition  of,  342 
Medulla  oblongata,  605,  606,  610 

as  a  conducting  path,  610 

connection    of,    with    cerebrum   and 
cerebellum,  6n 

functions  of,  612 

reflex  centers  of,  612 

tracts  through,  612 
Medullary  sheath,  66 
Medullated  fibers,  66 
Meissner's  corpuscles,  75 
Melanins,  96 
Membrana  decidua,  788 

propria,  23 

tympani,  701 

Membranous  labyrinth,  704 
Mendelian  inheritance  and  chromosomes, 

785 

Menstrual  discharge,  778 
Menstruation,  777 
Mesoblast,  u 
Mesothelium,  26 
Metabolism,  7,  454 

basal  rate,  478 

calories  per  square  meter  per  hour, 
478 

constructive,  7 

destructive,  7 

during  fasting,  472 

endogenous,  459 

exogenous,  459 

height,  weight  chart,  478 

influence  of  ductless  glands  on,  482 
of  reproductive  glands  on,  498 

intermediate,  459 

nutrition  and  diet,  454 

of  carbohydrates,  467 

of  fats,  463 

of  proteins,  458 
Metaphase,  20 
Meta  plasm,  17 
Metaproteins,  87,  95,  112 
Methemoglobin,  142 
Microcytes,  128 

Micro-organisms  in  intestines,  391 
Microsomes,  9 
Micturition,  440 

center  for,  597 
Mid-brain,  606,  623 


Milk,  composition  of,  343,  794 

sugar,  101 

Millon's  reaction,  107 
Mineral  foods,  341,  344,  471 

absorption  of,  in  intestines,  419 
Minimal  stimulus,  519 
Mitosis,  19 
Molisch  reaction,  108 
Monaster,  20 
Monosaccharides,  99 
Mossy  fibers,  619 
Motor  areas  of  cortex,  639,  641 
of  human  brain,  641 

end-plates,  63 

impulses,  602 

-oculi  nerve,  651 

tracts  in  human  brain,  643 
Mouth,  absorption  in,  412 

digestion  in,  347 

in  speech,  554 
Movement,  ameboid,  3 

gliding,  5 

streaming,  4 
Movements,  circus,  623 

forced,  623 

of  stomach,  374 
Mucigen,  353 
Mucinogen,  349 
Mucins,  95,  353 
Mucoids,  95,  in 
Mucous  membranes,  336 
Mucus  in  urine,  434 
Murexide  test,  432 
Muscle,  blood  supply  of,  63 

cardiac,  62,  195,  529 

chemical  changes  of,  516 
composition  of,  511 

coagulation  of,  510 

conditions    affecting    irritability    of, 

5i9 

conductivity  of,  513 
contractility  of,  512 
contraction  of,  512,  531,  560 

phase  of,  513 
contracture,  524 
currents,  demonstration  of,  516 
development  of,  59,  64,  65 
effect  of  blood  supply  on,  522 

of  drugs  on,  523 

of  nerve  supply,  523 

of  temperature  on,  521 

of  use  on,  520 


8io 


INDEX 


Muscle,  elasticity  of,  511 

electrical  phenomena  of,  516 

end-plates,  63 

experiments  on,  557 

ferments,  510 

in  rigor  mortis,  526 

involuntary,  58,  530,  569 

compared  with  skeletal  and  car- 
diac, 531 

irritability  of,  512,  519,  559 

-nerve  physiology,  510 
preparation,  558 

nerve  supply  of,  63 

non-striated,  58 

plain,  58 

plasma,  510 

properties  of,  511 

record  of  contraction  of,  521 

serum,  510 

skeletal,  60 

smooth,  531 

spindles,  78 

stimuli,  519 

striated,  60 

tetanus,  525,  565 

-tone,  center  of,  596 

voluntary,  60 

Muscular  action  as  heat  producer,  503 
•center  for  tone  of,  596 
contraction,  524 
action  currents,  516 
apparatus  for  producing  and  re- 
cording, 514 

changes  in  shape  during,  515 
characteristics  of  single,  513 
chemical  changes  during,  516 
conditions  affecting  character  of, 

579 

co-ordinated,  524 
differences  between  voluntary  and 

involuntary,  530 
effect  of  blood  supply  on,  522 
of  drugs  on,  523 
of  fatigue  on,  564,  561 
of  load  on,  562 
of  nerve  supply  on,  523 
of  rate  of  stimulation  on,  523 
of  repeated  activity  on,  520 
of  strength  of  stimulus  on,  519, 

560 
of    temperature    on,    521,    562, 

565 


Muscular     action,     effect    of    use     on, 

520 

electrical  changes  during,  516 
energy  liberated  during,  518 
heat  produced  during,  517 
latent  period   of,  514,  564 
metabolism  during,  528 
refractory  phase  of,  529 
response    to   stimuli  in   voluntary 

and  involuntary,  530 
simple,  564 

summation  of  contractions,  524 
tetanic,  523,  565 
voluntary,  523 
contracture,  524 
co-ordination,  524 
energy,  518 
sense,  681 
tissue,  58 
Musculi  papillares,  1 73 

pectinati,  169 
Mydriasis,  663 
Myelin  sheath,  66 
Myelocyte,  106,  116,  134 
Myeloplaxes,  42 

Myogenic  theory  of  heart  beat,  202 
Myogram,  513 
Myograph,  pendulum,  563 
Myohematin,  511 
Myopia,  737 
Myosin,  511 

ferment,  511 
Myosinogen,  511 
Myxedema,  485 

Near-point,  731,  760 
Nephrolytic  sera,  150 
Nerve  cells,  65,  572 

arrangement  in  spinal  cord,  582 

body,  72 

characteristic  of  individual,  575 

functions  of,  575 

in  cerebral  cortex,  527 

neurone  theory,  573 

nutritive  influence  of,  575 

transmission  of  impulses  through, 

577 

centers,  578 
collaterals,  69 
end-brushes,  70 
fibers,  65,  66 

effect  of  battery  current  on,  535 


INDEX 


Nerve  fibers,  fatigue  of,  575 
functions  of,  534 
medulla  ted,  66 
non-medullated,  67 
impulses,  533 
cellulifugal,  577 
cellulipetal,  577 
character  of,  533 
rate  of,  570 
specific  energy  of,  576 
transmission  through  neurone,  577 
velocity  of,  534 
stimuli,  512,  533 
terminations,  74 
Nerves,  cardiac,  214,  266 
cranial  functions  of,  650 
depressor,  248 

effect  of  currents  on  human,  535,  537 
experiments  on,  557 
irritability  of,  558 
vasoconstrictor,  246,  254 
vasodilator,  249,  251,  252 
vasomotor,  243,  246,  249 
Nervous  system,  572 

axones  of,  65,  577 
dendrites  of,  65,  577 
functions  of,  572 
ganglia  of,  578 

influence  on  secretion,  339,  446 
laboratory  experiments  on,  675 
neuroglia  of,  65,  78 
spinal  hemisection,,677 
sympathetic,  663 
Neurilemma,  66 
Neurofibrils,  72 

Neurogenic  theory  of  heart  beat,  201 
Neuroglia,  65,  78 
Neurone,  65,  572 

characteristics  of,  575 

irritability,  675 

polarity  in,  675 

theory-,  573 

transmission      of      nerve     impulses 

through,  577 
types  of,  578 
Neutrophiles,  116,  132 
Ninth  cranial  nerve,  659 
Nitrogen  in  proteins,  106 
Nitrogenous  equilibrium,  457 
food,  341 
output,  459 
substances  in  body,  79 


Nitrous  oxide,  effect  of  breathing,  318 
Nodes,  cardiac,  204 

of  Ranvier,  66 

Nostrils,  respiratory  movements  of,  291 
Nuclei  of  optic  thalamus,  625 
Nucleic  acid,  92 
Nucleoalbumins,  95 
Nucleoli,  1 8 
Nucleo proteins,  87,  92 
Nucleus,  9,  1 8 

ambiguus,  657 

ruber,  624 
Nutritional  diseases,  480 

Obesity,  467 
Ocular  fixation,  751 
Odontoblasts,  57 
Oils,  as  food,  344 
Olfactory  apparatus,  694 

bulb,  696 

center,  646 

glomeruli,  646 

nerve  and  tract,  646 
Olivary  bodies,  608,  610 
Olive,  accessory,  610 

superior,  616 
Onkograph,  438 
Onkometer,  437 
Ophthalmoscope,  742,  766 
Opsonic  index,  151 
Opsonins,  151 
Optic  center,  645 

nerve,  719 

thalami,  625 
Optical  apparatus,  724 

defects  in,  734 
Organ  of  Corti,  705 
Osmosis,  152 

Osmotic  pressure,  85,  153 
Osseous  labyrinth,  703 
Ossicles  of  ear,  701 
Ossification,  46 

center  of,  47 

in  membrane,  46 
Osteoblasts,  47 
Osteoclasts,  49 
Osteogenetic  fibers,  47 
Output  of  energy,  479 
Ovaries,  772 

relation  to  metabolism,  499 
Oviducts,  775 
Ovulation,  777 


812 


INDEX 


Ovum,  783 

changes  following  impregnation,  786 

prior  to  impregnation,  783 
Oxalic  acid  in  urine,  435 
Oxygen,  amount  excreted,  456 

determination  of,  in  air,  329 

in  blood,  302,  334 

in  expired  air,  298 

in  tissues,  308 

of  the  blood,  302 

Van  Slyke  apparatus,  301 
Oxyhemoglobin,  139,  164,  306 

Pacinian  corpuscles,  75,  379 
Pain,  sense  of,  68 1 
Pancreas,  377 

enzymes  of,  381 
extirpation  of,  492 
internal  secretion  of,  491 
islands  of  Langerhans  in,  379,  492 
secretion  of,  379 
structure  of,  377 
Pancreatic  digestion,  372,  406 

cleavage  products  of,  407 
fistula,  379 
glycemia,  470 
juice,  379,  406 
artificial,  407 
chemical  characters  of,  406 

composition  of,  379 
enzymes  of,  381,  407 
action  of,  381,  407 
conditions  influencing  action  of, 

383 
secretion  of,  406 

action  of  nerves  on,  380 
influence  of  secretin  on,  406 
Papillae  of  skin,  442 
of  tongue,  691 

Paralytic  secretion  of  saliva,  351 
Paramyosinogen,  511 
Parathyroidectomy,  487 
Parathyroid  glands,  487 
Parietal  association  center,  650 
Parotid  gland,  348 
nerves  of,  352 
Parturition,  792 
center,  598 

Peduncles  of  cerebrum,  623 
Pellagra,  481 

Pelvic  viscera,  vascular  nerves  for,  257 
Penis,  771 


Pepsin,  347,  370 

action  of,  370 
Pepsinogen,  370 
Peptides,  88,  96 
Peptone  plasma,  126 
Peptones,  88,  96,  112,  370 
Perforating  fibers  of  Sharpey,  46 
Pericardium,  167 
Perichondrium,  39,  47 
Perimysium,  60 
Perineurium,  69 
Periosteum,  43 
Peristalsis,  intestinal,  395 

of  stomach,  375 

reversed,  395 

Perivascular  lymphatic  sheaths,  1 79 
Perspiration,  445 
Pes,  623 

Pfluger's  law  of  contractions,  536 
Phagocytes,  158 
Phagocytosis,  158 
Phakoscope  of  Helmholtz,  762 
Phenomena  of  life,  i 
Phenomenon  of  treppe,  561 
Phenylalanine,  81 
Phosphates,  103,  115 
Phosphoproteins,  87,  95 
Phosphoric  acid  in  urine,  434,  451 
Phosphorus  in  proteins,  106 
Phrenic  nerve,  influence  on    respiration, 

332 
Pigments,  bile,  408 

in  urine,  433,  452 
Pituitary  body,  494 

action  of  grafts,  extracts,  etc.,  497 

effect  of  removal  of,  496 

function  of,  495 
Placenta,  788 

Plants  differentiated  from  animals,  12 
Plasma,  117,  136 

chemistry  of,  163 

composition  of,  136 
Plasmosomes,  18 
Plethysmogram,  277 
Plethysmograph,  245,  276 
Pleurae,  282 

Pneumogastric  nerve,  660;  and  see  Vagus. 
Pneumograph,  292 
Polysaccharides,  99 
Pons  Varolii,  605,  615 
Postdicrotic  wave,  239 
Posterior,  longitudinal  bundle,  616 


INDEX 


8i3 


Posterior,  pyramids,  606 

roots  of  spinal  nerves,  489,  594 
Postganglionic  fibers,  666 
Potassium  salts  in  the  body,  103,  104 
Poultry,  composition  of,  343 
Precipitins,  151 
Predicrotic  wave,  239 
Preganglionic  nerve  fibers,  666 
Presbyopia,  739 
Pressor  nerves,  249 
Pressure,  endocardiac,  187 
Pronucleus,  female,  784 

male,  787 

Prostate  gland,  771 
Protamines,  87,  91 
Proteins,  80 

absorption  of,  from  intestines,  416 

action  of  trypsin  on,  381 

as  fat  formers,  459 

as  foods,  341 

as  glycogen  formers,  459 

circulating,  458 

classification  of,  86,  88 

coagulation,  109 

color  reactions  of,  107 

crystals,  105 

decomposition  products,  381 

derived,  87,  95,  112 

digestion  of,  369,  381 

experiments  on,  105,  109 

hydrolytic  cleavage  of,  83,  84 

metabolism  of,  458 

nitrogen  and  phosphorus  in,  106 

precipitation^  reactions  of,  108 

preparation  of,  105 

properties  of,  85 

reactions  of,  107 

salting  out  experiments  on,  109 

simple,  86 

sulphur  in,  106 

tissue,  458 

wheat,  83 

Proteolytk  ferments,  347 
Proteoses,  88,  96,  112,  370 
Prothrombin,  124 
Protoplasm,  i,  2 

chemistry  of,  3 

differentiation  of,  8,  10 

growth  of,  7 

irritability  of,  5 

metabolism  in,  7 

movement  of,  3 


Protoplasm,  physiological  characteristics 

of,  3 

reproduction  of,  8 
Ptosis,  652 

Ptyalin,  action  of,  347,  355,  402 
Pulse,  236 

arterial,  270,  272 

dicrotic,  240 

variations  in  rate  of,  212 

-wave,  rate  of  propagation  of,  272 
Pulvinar,  626 
Pupil,  718 

contraction  of,  733 

dilatation  of,  733 
center  for,  614 
Purines,  94 
Purkinje's  cells,  618,  620 

fibers,  171,  203 

figures,  740 

shadows,  764 

Purkinje-Sanson's  images,  762 
Putamen,  627 

Putrefaction  in  intestines,  399 
Pyloric  glands,  364 

orifice,  374 
Pyramids,  606 

decussation  of,  607 
Pyramidines,  93 

Racemose  glands,  337 
Ranvier,  nodes  of,  66 
Reaction  of  degeneration,  539 
Red  corpuscles,  126 

action  of  reagents  on,  157 
chemical  composition  of,  139 
development  of,  129 
origin  of,  129 
Red  nucleus,  624 
Reflex  action,  590 
time  of,  594 
arc,  590 

centers  in  medulla,  612 
Reflexes,  complex,  592 
cutaneous,  599 
inhibition  of,  598 
muscle,  599 
simple,  592 

special  centers  for,  596 
spinal,  595 
Refraction,  759 
Refractory  period,  199 
phase,  529 


8i4 


INDEX 


Relaxation  phase  of  muscle,  514 
Remak's  fibers,  67 

ganglia,  196 
Rennin,  347,  373 

action  of,  373,  406 
Reproductive     glands,     relation     of     to 

metabolism,  498 
organs,  768 
of  female,  772 
of  male,  768 
Reserve  air,  295,  328 
Residual  air,  295 
Respiration,  278 

changes  in  diameter  of  chest,  dur- 
ing, 327 

effect  of  altitude  on,  318 
effect  on  circulation,  323 
of  atmospheric  pressure  on,  318 
of  various  gases  on,  318 
of  vitiated  air  on,  317 
expiration,  290 

influence     of     cutaneous     nerves 

on,  331 
of  general  sensory  nerves  on,  313, 

33i 
of    glosso-pharyngeal    nerves    on, 

3i3 

of  phrenic  nerves  on,  332 

of   superior   laryngeal   nerves   on, 
312 

of  vagus  nerves  on,  311,  331 
inspiration,  287 
internal,  279 

laboratory  experiments  in,  327 
mechanism  of,  287 
nervous  apparatus  of,  310,  331 
rate,  327 
rhythm  of,  291 
special  types  of,  315 
tissue,  279 

volume  of  air  breathed,  299,  328 
Respiratory  apparatus,  279 

elimination  of  carbon  dioxide  by, 

309 

nervous  regulation  of,  310 
capacity,  295,  328 

circumstances  affecting,  296 
center,  310,  614 

automatic  action  of,  313 

stimulation  of,  313 
changes  in  air  breathed,  297 

in  the  blood,  301 


Respiratory  changes  in  the  tissues,  308 

interchange,  330 

movements,  287 
character  of,  326 
establishment  of,  at  birth,  315 
nervous  mechanism  of,  331 
of  nostrils  and  glottis,  291 
rate  and  character  of,  331 
recording  of,  291 
relative  time  of,  293 

murmur,  294 

muscles,  force  of,  297 

pressure,  301,  328 

quotient,  300 

rate,  296,  327 

rhythm,  293 

action  of  stimuli  on,  311 

terms  for  quantity  of  air  breathed, 

294 
Resuscitation  from  drowning,  321 

from  electric  shock,  321 
Rete  mucosum,  442 
Reticular  formation  in  medulla,  608 
Reticulum,  9,  17 
Retina,  719 

cones  of,  721 

inverted  image  on,  760 

layers  of,  720 

localization  in,  744 

movements  of  pigment  cells,  746 

rods  of,  721 
Retinal  image,  duration  of,  765 

relation  of  size  to  distance,  764 
Retinoscopy,  766 
Rheoscopic  frog,  534 
Rhodopsin,  745 

Rhythmical  contractility  of  heart,  196 
Rhythmicity  of  arterial  flow,  230 
Ribs,  movement  of,  in  respiration,  287 
Rigor  mortis,  526 

heart,  180 

heat,  527 

order  of  occurrence,  527 
Rima  glottidis,  279 
Rolandic  area,  642 
Running,  543 

Saccharose,  100 
Sacculus,  704,  715 
Sacral  autonomies,  671 
Saliva,  354 

action  of,  on  starch,  355,  356,  401 


INDEX 


8iS 


Saliva,  chemical  composition  of,  354 
function  of,  355 
properties  of,  355 
ptyalin  in,  355 
quantity  of,  354 
secretion  of,  center  for,  349 
nerve  mechanism  of,  349 
rate  of,  354 
Salivary  digestion  in  stomach,  358 

influence  of  acids  and  alkalies  on, 

401 

of  temperature  on,  401 
glands,  348 

changes  in  during  secretion,  352, 

400 

nerves  of,  399 
structure  of,  348 
secretion,  352 
reflex,  399 

Salting  out  proteins   109 
Salts,  absorption  of,  by  intestines,  419 
as  foods,  344 
bile,  386,  408 
in  the  body,  103 

tests  for,  115 
Sanson's  images,  730 
Saponification,  114,  382 
Sarcode.  2 
Sarcolemma,  61 
Sarcoplasm,  62 
Sarcostyles,  61 

Sarcous  elements  of  Bowman,  61 
Schaefer's  method  of  resuscitation,  322 
Schemer's  experiment,  760 
Schwann,  sheath  of,  66 
Scurvy  and  vitamine,  C,  481 
Sebaceous  glands,  444 
Secretin,  390 

influence    on    pancreatic    secretion, 

390,  406 
Secreting  glands,  335 

production  of  heat  by,  503 
types  of,  336 
Secretion,  390 

circumstances  influencing,  339 
discharge  of,  339 
external,  335 
internal,  335,  482 
organs  and  tissues  of,  336 
process  of,  338 
psychic,  403 
Segmentation,  787 


Semicircular  canals,  703,  714 
Semilunar  valves,  183 

action  of,  183 
Seminal  fluid,  771 

vesicles,  772 
Sensations,  binaural,  712 

common,  679 

objective,  680 

of  color,  746 

special,  680 

subjective,  680 
Sense,  hearing,  699,  706 

muscular,  687 

of  equilibrium,  713 

of  pain,  686 

of  sight,  715 

of  smell,  694,  757 

of  taste,  659,  689,  757 

of  temperature,  685,  756 

of  touch,  68 1 

organs,     directions    for     experiment 
on,  756 

perceptions,  68 1 
Senses,  the,  679 

internal,  679 

special,  680 
Sensorium,  680 
Sensory  areas  of  brain,  644 

illusions,  680 

impulses,  60 1 
Serine,  81 

Serous  membranes,  336 
Serum,  118,  135 

agglutinative  substances,  150 

blood,  1 1 8,  135 

chemistry  of,  163 

composition  of,  136 

globulicidal  action  of,  149 

hemolytic  action  of,  149 

muscle,  510 

precipitins  of,  151 
Seventh  cranial  nerve,  657 
Sex,  germ  cells  and  determination  of,  786 
Sex-linked  inheritance,  786 
Sharpey's  fibers,  46 
Sight,  715 
Silicon,  104 

Sino-auricular  node,  204 
Sinus  arrhythmia,  213 
Sixth  cranial  nerve,  656 
Size,  estimation  of,  689,  755 
Skin,  absorption  from, "420 


8i6 


INDEX 


Skin,  carbon  dioxide  exhaled  by,  446 

excretion  by,  444,  456 

excretory  function  of,  441,  444 

functions  of,  441,  444 

glands  of,  424 

loss  of  heat  from,  504 

structure  of,  441 

water  excreted  by,  445 
Sleep,  673 
Small  intestine,  absorption  in,  414 

digestion  in,  390 
Smell,  center  for,  646 

sensation  of,  757 

sense  of,  694 

Sodium  salts  in  the  body,  103,  104 
Solidity,  judgment  of,  754 
Solutions,  isotonic,  154 
Somesthetic  area  of  brain,  644 
Somnambulism,  674 
Sound,  707 
Sounds,  articulate,  555 

localization  of,  711 

of  the  heart,  183 

pitch  of,  707 
Speech,  544,  554 

action  of  tongue  in,  555 

of  mouth  in,  555 
Spermatids,  782 
Spermatocytes,  781 
Spermatogonia,  781 
Spermatozoa,  783,  785,  786 
Spherical  aberration,  734,  761 
Sphygmogram,  239 
Sphygmograph,  238 
Sphygmomanometer,  226,  239 
Sphygmometer,  238 
Spinal  accessory  nerve,  662 

centers,  596,  597,  598 

cord,  580 

anterior  pyramidal  tract,  586 
antero-lateral     descending      tract, 

587 

arrangement  of  nerve  cells  in,  582 
ascending  degeneration,  tracts  of, 

587 

comma  tract  of,  587 
conduction  in,  600 
course  of  motor  impulses  in,  602 

of  sensory  impulses  in,  601 
descending     degeneration,     tracts 

of,  586 
direct  cerebellar  tract,  587 


Spinal  cord,  fasciculi  of,  584 
functions  of,  590 
general  features  of,  580 
Gowers'  tract,  562 
irradiation  of  impulses  in,  593 
lateral  pyramidal  tract,  586 
peculiarities   of   different    regions, 

590 

reflex  action  in,  590 
tracts  of,  584 
weight  of,  632 

nerve-roots,  functions  of,  588 
nerves,  588 

anterior  roots,  588 
course  of  fibers,  588 
posterior  roots,  589 
reflexes,  595 
Spirem,  20 
Spirometer,  295 

Spleen,  vascular  nerves  for,  257 
Spongioplasm,  9,  17 
Staircase  contractions,  520 
Stammering,  556 
Starch,  101 

action  of  amylopsin  on,  382 

of  ptyalin  on,  356 
animal,  102 

chemical  reactions  on,  113 
hydrolysis  of,  113 
Starvation,  472 

death  from,  474 

effect  on  body  temperature,  473 
influence  of,  on  excretions,  474 
symptoms  of,  473 
Steapsin,  347,  382 
Stercobilin,  387 
Stereoscope,  754 
Stethograph,  292 
Stimuli,  forms  of,  512 

maximal  and  minimal,  510,  560 
Stokes'  fluid,  141 
Stomach,  360 

absorption  from,  412 

action  of  pylorus,  374 

blood-vessels  of,  364 

changes  in  glands  during  secretion, 

365 

digestion  in,  358,  360,  372 
gases  in,  394 
glands  of,  363,  365 
hunger  contraction  of,  375 
lymphatics  of,  364 


INDEX 


8l7 


Stomach,  movements  of,  373 

nerves  of,  360 

pancreatic  digestion  in,  372 

peristalsis  of,  374 

secretion  in,  365 

structure  of,  361 

vascular  nerves  for,  257 
Stomata,  26 
Stratum  granulosum,  442 

lucidum,  442 

Malpighii,  442 
Striated  muscle,  60 

development  of,  64 
Sublingual  gland,  348 
Submaxillary  gland,  348 

action  of  atropine  on,  351 

influence  of  nerves  on,  350 

paralytic  secretion  of,  351 

secretion  of,  354 
Substantia  nigra,  623 
Succus  entericus,  390 
Sucking,  center  for,  614 
Sudoriferous  glands,  443 
Sugar,  test  for,  in  urine,  453 
Sulphates  in  body,  115 
Sulphur  in  proteins,  106 
Sulphureted  hydrogen,  effect  of  breath- 
ing, 318 

Sulphuric  acid  in  urine,  431,  434,  450 
Sulphurous    acid,     effect    of    breathing, 

.3l8 
Summation,  524 

of  stimuli,  592 
Superior  laryngeal  nerve  in  respiration, 

309 

Supplemental,  air,  295 
Suprarenal  capsules,  488 
functions  of,  489 
internal  secretion,  490 
nerves  of,  491 
Swallowing,  352 
Sweat,  445 

centers,  615 

chemical  composition  of,  445 
glands,  443 

influence  of  nervous  system  on  se- 
cretion of,  446,  449 

Sympathetic  ganglia,  functions  and  struc- 
ture, 663,  666,  668 
afferent  fibers  in,  672 
functions,  670 
Synapse,  591 


Synovial  membranes,  663 
Systole  of  heart,  1 79 

duration  of,  181 

extra  ventricular,  209 

Tactile  corpuscles,  76,  683 

of  Meissner,  75 
menisques,  77 
Taste,  689 

acuteness  of,  691 
after-,  693 
buds,  280,  690 
center,  693 
contrasts,  693 

influence  of  fifth  nerve  on,  655 
nerves  of,  689 
seat  of,  693 
sensation  of,  692,  757 
sense  of,  689 
varieties  of,  692 
Teeth,  51 

dentine  of,  53 
development  of,  56 
enamel  of,  54 
ivory  of,  53 
permanent,  52 
structure  of,  52 
temporary,  51 
Tegmentum,  616 
Telophase,  21 
Temperature,  body,  503 
dissipation  of,  501 
influence    of    extreme    heat    and 

cold  on,  505 
of  starvation  on,  472 
regulation  of,  503 
sense  of,  685,  756 
variations  in,  501 

influence   of,   on    muscular   contrac- 
tion, 565 

Tenth  cranial  nerve,  660 
Testes,  768 

relation  to  metabolism,  499 
Tetanometer,  565 
Tetanus,  524,  565 
Thalami,  625 
Thermogenic  centers,  509 
Third  cranial  nerve,  651 
Thoracic    viscera,    vascular    nerves    for, 

257 

Thoracic  autonomies,  669 
Thoracograph,  327 


8i8 


INDEX 


Thorax,    respiratory   changes   in    diame- 
ter, 289,  327 
Thrombin,  121 
Thrombocytes,  135 
Thrombogen,  122 
Thrombokinase,  122,  347 
Thymine,  94 
Thymus  gland,  494 
Thyroidectomy,  485 
Thyroid  feeding,  485 

gland,  484 

accessory,  484 
functions  of,  484 

grafting,  485 

hormone,  486 
Thyroiodine,  486 
Tidal  air,  282,  328 
Tissues,  connective,  33 

elementary,  23 

epithelial,  23 

interchange  of  gases  in,  308 

lymphoid,  36 

muscular,  58 

nervous,  65,  67 
Tone,  of  artery,  244 

of  muscle,  596 
Tongue,  685 

action  of,  in  speech,  556 

papillae  of,  683 

Tonicity  of  heart  muscle,  198 
Tonometer,  304 
Tooth-pulp,  52 
Touch  corpuscles,  75,  76,  683 

sense  of,  68 1 

acuteness  of,  683 
Trabeculae  carneae,  178 
Trachea,  280 
Tract  of  Burdach,  587 

of  Goll,  587 

of  Cowers,  587 
Traube-Hering  curves,  249 
Treppe,  phenomenon  of,  561 
Tricuspid  valve,  action  of,  181 
Trigeminal  nerve,  653 
Tri-olein,  97 
Tri-palmitin,  97 
Tri-stearin,  97 
Trochlearis  nerve,  653 
Trunk,  vascular  nerves  for,  258 
Trypsin,  379,  381 

action  of,  381 
Tryptophane,  82 


Tubular  glands,  337 
Tubuli  seminiferi,  768 

uriniferi,  421 

Twelfth  cranial  nerve,  663 
Tyco  sphygmomanometer,  226 
Tympanum,  701 
Tyrosine,  81 

Unorganized     ferments,     346;     and     see 

Enzymes. 

Unstriped  muscle,  58 
Uracil,  94 
Urea,  amount  in  tissues,  431 

antecedents  of,  462 

determination  of,  451 

formation  of,  431,  442 

preparation  of,  451 

properties  of,  430 

quantity  excreted,  432 
Ureters,  427 
Uric  acid,  94,  432,  451 

condition  of,  in  urine,  432 

formation  of,  463 

properties  of,  432 

tests  for,  452 
Urinary  bladder,  427 
Urine,  427 

abnormal  constituents  of,  452 

albumin  in,  435,  453 

ammonia  in,  433 

analysis  of,  449 

average     daily     quantity     of     con- 
stituents, 429 

chlorides  in,  435,  450 

composition  of,  428 

creatinin  in,  433,  452 

cystin  in,  435 

dextrose  in,  435 

discharge  of,  440 

diuretics,  action  of,  439 

excretion  by,  455,  456 
experiments  on,  437 

factors  affecting  secretion  of,  436 

general  properties  of,  427 

hippuric  acid  in,  433,  463 

indican  in,  452 

mucus  in,  434 

nitrogenous  substances  in,  430,  452 

occasional  constituents  of,  435 

oxalic  acid  in,  435 

phosphates  in,  434,  451 

pigments  in,  433,  452 


INDEX 


819 


Urine,  quantity  of,  429,  449 

reaction  of,  428,  449 

relation  of  blood  pressure  to  secre- 
tion of,  448 

saline  matter  in,  434 

secretion  of,  theories  of,  436 

solids  of,  450 

specific  gravity  of,  428,  449 

sugar  in,  453 

sulphates  in,  434,  450 

urea  in,  430,  431,  451 

uric  acid  in,  410,  432 

variations   in   quantity  of   constitu- 
ents, 430 

in  specific  gravity,  429 
Uriniferous  tubules,  422 
Urobilin,  433,  452 
Urochrome,  433 
Uroerythrin,  434 
Uterine  tubes,  775 
Uterus,  776 
Utriculus,  704,  715 

Vagina,  776 
Vagus  nerve,  660 

distribution  of,  660 

effects  of  section,  662 

functions  of,  662 

relation  to  deglutition,  660 
to  gastric  secretion,  662 
to  heart's  action,  662 
to  respiration,  312,  332 
Valine,  82 

Valves  of  heart,  169 
action  of,  181 

of  veins,  178 

Van  Slyke  blood  gas  apparatus,  303 
Vasoconstrictor  activity,  250 

center,  245 

nerves,  244,  246,  252,  259 

reflexes,  247 
Vasodilator  activity,  250 

center,  250 

nerves,  249,  252 

reflexes,  250 
Vasomotor  centers,  247 

changes,  275 

nerves,  243 

tone,  246 
Veins,  177 

blood  pressure  in,  227 

structure  of,  178 


Veins,  valves  of,  178 

vasoconstrictor  nerves  in,  259 
Venous  blood,  analysis  of,  302 

flow,  234 

velocity  of,  235 
Ventilation,  317 

Ventricles  of  heart,  action  of,  180 
Vesico-spinal  center,  597 
Vesiculae  seminales,  771 
Vesicular  breathing,  294 
Vicq  d'  Azyr,  bun-die  of,  625 
Villi,  414 

Visceral  sensations,  60 1 
Vision,  accommodation  of,  728,  731 

binocular,  751 

field  of,  744 

limits  of,  760,  765 

localization  of,  744 

mechanism  of  accommodation,  731 

range  of  distinct,  731 
Visual  acuity,  767 

center,  645 

image,  projection  of,  732 

judgments,  754 

purple,  745 

sensations,  739 
after-images,  741 
duration  of,  741 
•  sense,  715 
Vital  capacity,  313,  328 

phenomena,  i 
Vitamine  A,  481 
Vitamine  B,  480 
Vitamine  C,  481 
Vitamines,  480 
Vitiated  air,  effects  of,  317 
Vocal  cords,  279,  545 

movements  of,  551 
Vocalization,  551 
Voice,  544 

difference     between     male    and     fe- 
male, 553 

production  of,  544 

quality  of,  554 

in  singing,  and  speaking,  552 

vocal  range  of,  551 
Vomiting,  376 

center  for,  376 

nervous  mechanism  of,  376 
Vowels,  555 

Walking,  539,  541 


820  INDEX 

Wallerian  degeneration,  575,  586  White    fibrous    tissue,    development    of, 
Water,  104  37 

absorption  of,  in  intestines,  419 

amount  excreted,  445,  455,  456  Xanthine,  94 

in  expired  air,  300  Xantho-proteic  reaction,  107 
of  the  body,  104. 

as  food,  345  Yellow  elastic  tissue,  35 
rigor,  526  chemical  composition  of,  91 

Weights  and  measures,  tables  of,  796  development  of,  37 

White  fibrous  tissue,  34 

chemical  composition  of,  90  Zymogens,  378 


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30m-10,'61  (C3941s4)4128 


194801 


